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REVUE SUISSE
DE
ZOOLOGIE
REVUE SUISSE
ZOOLOGIE
ANNALES
DE LA
SOCIETE SUISSE DE ZOOLOGIE
ET DU
MUSEUM D'HISTOIRE NATURELLE
DE LA VILLE DE GENEVE
la
GENEVE
2000
ISSN 0035-418X
TABLE DES MATIERES
TOME 107 — 2000
Fascicule 1
JUVARA-BALS, Ilinca & Vojciech WITALINSKI. Description of five new species of
Holoparasitus s. str. with redescription of H. apenninorum (Berlese, 1906)
and H. cultriger (Berlese, 1906) from Italy and Spain (Acari, Gamasida,
Barasıldao) Re AN RI DEA N ikl ee LR URI
SILVA, Carla Maria Menegola da & Beatriz MOTHES. Three new species of Geodia
Lamarck, 1815 (Porifera, Demospongiae) from the bathyal depths off
Brazilianicoast.s outhwestem- AtlantiGe EER a een
MAHUNKA, Sandor & Luise MAHUNKA-PAPP. Oribatids from Switzerland III
(Acari: Oribatida: Oppiidae | and Quadroppiidae). (Acarologica Genavensia
GIL DE PERTIERRA, Alicia A. & Alain de CHAMBRIER. Rudolphiella szidati sp. n.
(Proteocephalidea: Monticelliidae, Rudolphiellinae) parasite of Luciopi-
melodus pati (Valenciennes, 1840) (Pisces: Pimelodidae) from Argentina
with new observations on Rudolphiella lobosa (Riggenbach, 1895)........
BENJAMIN, Suresh P. & Rudy JOCQUÉ. Two new species of the genus Suffasia
Pomorie wanka (Araneae: ZOd arid ae) se een me et ee ee en ee) stat IE
LANG, Claude. Diversité du zoobenthos dans 47 rivières du canton de Vaud:
tendances ES 1 O ieee ee VOIR En
Vit, Stanislav. Contribution a la connaissance de la famille Eucinetidae (Cole-
Oi OUST) ER N SR E PE Er IE a a RR NEE
BANZIGER, Ruth. Spatio-temporal distribution of size classes and larval instars of
aquatic insects (Ephemeroptera, Trichoptera and Leopidoptera) in a Potamo-
ecton pecinatus ds bed (Make'Geneva, Switzerland)... 2.20... RENE
BESUCHET, Claude & Stanislav Vit. Les Nanophthalmus Motschulsky d’Europe
(@oleoptera,seydmaenidae) ZERI
Heyp, Andreas & Wolfgang PFEIFFER. Uber die Lauterzeugung der Welse (Silu-
roidei, Ostariophysi, Teleostei) und ihren Zusammenhang mit der Phylo-
sentetundsderSchreekreaktions rn A ey eae
GANTENBEIN, Benjamin, Christian KRoPF, Carlo Rodolfo LARGIADER & Adolf
SCHOLL. Molecular and morphological evidence for the presence of a new
Buthid taxon (Scorpiones: Buthidae) on the Island of Cyprus.............
Fascicule 2
LANG, Claude. Etat trophique du lac de Morat indiqué par le zoobenthos: tendance
LO SOB OS eae Maia arenes as nr NEST ae Se nO
AZPELICUETA, Maria de la Mercedes & José O. GARCIA. A new species of Astya-
nax (Characiformes, Characidae) from Uruguay river basin in Argentina,
withrremarkssonthooksptesenceiin@haracıdaege nn. m ar
Pages
3-30
31-48
49-79
81-95
97-106
107-122
123-138
139-151
153-163
165-211
213-232
233-243
245-257
VI TABLE DES MATIERES
BALLERIO, Alberto. Revision of the genus Ebbrittoniella Martinez (Coleoptera:
Scarabacoideay Ceratocanthidae) 4... RE
LIENHARD, Charles. A new desert psocid from Namibia (Insecta: Psocoptera:
NiO SUN AS) Bene ee ise ists dle: ee ee eee ee
LEISTIKOW, Andreas. Terrestrial Isopoda from Guatemala and Mexico (Crustacea:
OniscideaxErimocheta eye se eee eee Cee eee
BURCKHARDT, Daniel & Mariängela GUAJARA. Euphalerus clitoriae sp. n., a new
psyllid species from Clitoria fairchildiana (Fabaceae, Papilionoideae), and
notes on other Euphalerus spp. (Hemiptera, Psylloidea).................
Koss, Kerstin & Wolfgang PFEIFFER. Zur Biologie des “Eigenweidefisches”
Carapus acus (Briinnich, 1768) (Carapidae, Teleostei), mit Hinweisen auf
einennicht-parasitische’Ernährung® PEER RR eee
BEDOS, Anne & Louis DEHARVENG. Un nouveau Collembole Neanurinae du Sud
du Vietnam, Blasconura batai sp. n., avec une clé des especes du genre
(@ollembolaxNeanurrdao)e NS ee E TORE DA o REA
BAUR, Hannes & Felix AMIET. Die Leucospidae (Hymenoptera: Chalcidoidea) der
Schweiz, mit einem Bestimmungsschlüssel und Daten zu den europäischen
TANDY, Mills & Jean-Luc PERRET. The Bufo tuberosus species group with the
description of a new species from the rainforest of Cöte-d’Ivoire..........
Fascicule 3
PUTHZ, Volker. The genus Dianous Leach in China (Coleoptera, Staphylinidae).
2OIAEontibutionitoitheiknowledse.of Steninae. 222. a. er sar ee
GATTOLLIAT, J.-L. & M. SARTORI. Contribution to the systematics of the genus
Dabulamanzia (Ephemeroptera: Baetidae) in Madagascar. ...............
MUSTER, Christoph & Konrad THALER. Das Männchen von Zelotes zellensis
Gurmmı(Araneae:/Gnaphosidae) er. LELE
VOGEL, Peter, Mark LAWRENCE & Ali AGHNAJ. Note sur les musaraignes
(Soricidae, Mammalia) du Pare National du Souss-Massa, Maroc..........
Lost, Ivan. A review of the Scaphidiinae (Coleoptera: Staphylinidae) of the
ReoplesssRepubliciof@hina; Mer IT aoe eee
Fascicule 4
POMORSKI, Romuald J. & Dariusz SKARZYNSKI. Redescription of Hymenaphorura
alticola (Bagnall, 1935) from the Alps and description of a new related
species from the Sudetes, Hymenaphorura improvisa sp. n. (Collembola:
OnychiunGae) i. lie Sa su. be oe ss dug +s) ae eee
ZAWADZKI, Claudio H., Roberto E. Reis & Erasmo RENESTO. Allozyme discri-
mination of three species of Loricariichthys (Siluriformes: Loricariidae) from
SouMenM Brazile dg. (000 NORTE
Pages
259-275
277-281
283-323
325-334
335-349
351-357
359-388
389-418
419-559
561-577
579-589
591-599
601-656
657-662
663-674
TABLE DES MATIERES
MAHUNKA, Sandor. Oribatids from Sabah (East Malaysia) VIII (Acari: Oribatida:
Dampfiellidae and Otocepheidae). (Acarologica Genavensia LXXXVI)......
LOURENÇO, Wilson R. More about the Buthoidea of Madagascar, with special
references to the genus Tiryobuthus Pocock (Scorpiones, Buthidae).........
PAGES, Jean. Japygidés (Diplura) du Sud-Est asiatique n° 9. - Dicellurata Gena-
DENSO PROIETTI REA EA a ASE ER Ne Le
COMELLINI, André. Notes sur les Psélaphines néotropicaux (Coleoptera, Staphy-
linidae, Pselaphinae) — 11. Un nouveau genre et sept espèces nouvelles de la
GRIDUEGESYMIELOPIASIM PE SE an PAUL ee ee
BÖHME, Wolfgang, Andreas SCHMITZ & Thomas ZIEGLER. A review of the West
African skink genus Cophoscincopus Mertens (Reptilia: Scincidae: Lygoso-
minae): resurrection of C. simulans (Vaillant, 1884) and description of a new
SPECIES IE LE ee N ee
ROWELL, C. H. F. Review of the Lithoscirtus genus group (Orthoptera, Acrididae,
Rroctolabmaec)) with description of mew species: ... 2.0... 00 ee ve eee
VALLAN, Denis. A new species of the genus Stumpffia (Amphibia: Anura:
Microhylidae) from a small forest remnant of the central high plateau of
Madagascar Sewerage cit Reese le scoot tua RN ce co ER
GANTENBEIN, Benjamin, Victor FET, Mark BARKER & Adolf SCHOLL. Nuclear and
mitochondrial markers reveal the existence of two parapatric scorpion species
in the Alps: Euscorpius germanus (C. L. Koch, 1837) and E. alpha
Caponaccoy 195 04statenov. (Euscorpiidac)ia- ea ale
LIENHARD, Charles. A new genus of Prionoglarididae from a Namibian cave
(insectasPSOCoptera) se ar. NE EN <a NE WB REN eo RE
MERZ, Bernhard. Two new Tephritidae (Diptera) from the Western Palaearctic
ICROM les SE cet SRR TOI OT NE ate EEE ER N ar
VII
Pages
675-720
721-736
737-764
765-776
777-791
793-834
835-841
843-869
871-882
883-892
DR
N
Lu
INDEX DES AUTEURS
par
ORDRE ALPHABETIQUE
AZPELICUETA, Maria de la Mercedes & José O. GARCIA. A new species of Astya-
nax (Characiformes, Characidae) from Uruguay river basin in Argentina,
with remarks on hook presence in Characidae. ........................
BALLERIO, Alberto. Revision of the genus Ebbrittoniella Martinez (Coleoptera:
Scarabacoideas Cerato Cani dI) EROI ee:
BANZIGER, Ruth. Spatio-temporal distribution of size classes and larval instars of
aquatic insects (Ephemeroptera, Trichoptera and Leopidoptera) in a Potamo-
geton pectinatus L. bed (Lake Geneva, Switzerland). ...................
BAUR, Hannes & Felix AMIET. Die Leucospidae (Hymenoptera: Chalcidoidea) der
Schweiz, mit einem Bestimmungsschliissel und Daten zu den europäischen
BEDOS, Anne & Louis DEHARVENG. Un nouveau Collembole Neanurinae du Sud
du Vietnam, Blasconura batai sp. n., avec une clé des especes du genre
(GollembolasiNeanunid ae) yes ee oes PR eine
BENJAMIN, Suresh P. & Rudy JOCQUÉ. Two new species of the genus Suffasia
fromesnilcankal(Aranecae=Zodarlidae) I re alle eect tie INDI one
‘ BESUCHET, Claude & Stanislav VIT. Les Nanophthalmus Motschulsky d’Europe
(ColeopteratScydmacnidae)y et. acess ese n
BÖHME, Wolfgang, Andreas SCHMITZ & Thomas ZIEGLER. A review of the West
African skink genus Cophoscincopus Mertens (Reptilia: Scincidae: Lygoso-
minae): resurrection of C. simulans (Vaillant, 1884) and description of a new
SPECIES See a STIRIA a Re ap OC AE MNT Al Ra AE
BURCKHARDT, Daniel & Mariangela GUAJARA. Euphalerus clitoriae sp. n., a new
psyllid species from Clitoria fairchildiana (Fabaceae, Papilionoideae), and
notes on other Euphalerus spp. (Hemiptera, Psylloidea). ................
COMELLINI, André. Notes sur les Psélaphines néotropicaux (Coleoptera, Staphy-
linidae, Pselaphinae) — 11. Un nouveau genre et sept espèces nouvelles de la
teibusdessMetopiasını 1.05 ar. EE IO TI
GANTENBEIN, Benjamin, Christian Kropr, Carlo Rodolfo LARGIADER & Adolf
SCHOLL. Molecular and morphological evidence for the presence of a new
Buthid taxon (Scorpiones: Buthidae) on the Island of Cyprus.............
GANTENBEIN, Benjamin, Victor Fer, Mark BARKER & Adolf SCHOLL. Nuclear and
mitochondrial markers reveal the existence of two parapatric scorpion species
in the Alps: Euscorpius germanus (C. L. Koch, 1837) and E. alpha
Caponiaccoml 950 statenovec(2usconpidae) a) er II ER
GATTOLLIAT, J.-L. & M. SARTORI. Contribution to the systematics of the genus
Dabulamanzia (Ephemeroptera: Baetidae) in Madagascar...............
245-257
259-275
139-151
359-388
351-357
97-106
153-163
777-791
325-334
765-776
843-869
561-577
x INDEX DES AUTEURS
GIL DE PERTIERRA, Alicia A. & Alain de CHAMBRIER. Rudolphiella szidati sp. n.
(Proteocephalidea: Monticelliidae, Rudolphiellinae) parasite of Luciopi-
melodus pati (Valenciennes, 1840) (Pisces: Pimelodidae) from Argentina
with new observations on Rudolphiella lobosa (Riggenbach, 1895)........
Heyp, Andreas & Wolfgang PFEIFFER. Über die Lauterzeugung der Welse (Silu-
roidei, Ostariophysi, Teleostei) und ihren Zusammenhang mit der Phylo-
genieund: der Schreckreaktion. sto. IRE OS PERE
JUVARA-BALS, Ilinca & Vojciech WITALINSKI. Description of five new species of
Holoparasitus s. str. with redescription of H. apenninorum (Berlese, 1906)
and H. cultriger (Berlese, 1906) from Italy and Spain (Acari, Gamasida,
Barasiudao)e E LORIA RI
KLoss, Kerstin & Wolfgang PFEIFFER. Zur Biologie des “Eigenweidefisches”
Carapus acus (Briinnich, 1768) (Carapidae, Teleostei), mit Hinweisen auf
einemicht-parasitische Ernährung... LO RE
LANG, Claude. Diversité du zoobenthos dans 47 rivières du canton de Vaud:
tendaneesl 98 NIIT, Ve ee A EE RS
LANG, Claude. Etat trophique du lac de Morat indiqué par le zoobenthos: tendance
OS FOO STE An RE i MR en ee
LEISTIKOW, Andreas. Terrestrial Isopoda from Guatemala and Mexico (Crustacea:
OniscideasGninocheta) a. Sr sae oe ET oe RI
LIENHARD, Charles. A new desert psocid from Namibia (Insecta: Psocoptera:
Miro sida) arg... sini We Seen ee news a, + CORE
LIENHARD, Charles. A new genus of Prionoglarididae from a Namibian cave
(InsectaaBsocoptera) at ila e die CES RE
LößL, Ivan. A review of the Scaphidiinae (Coleoptera: Staphylinidae) of the
ReoplessiRepubliciof China IE RER ERI RR
LOURENÇO, Wilson R. More about the Buthoidea of Madagascar, with special
references to the genus Tiryobuthus Pocock (Scorpiones, Buthidae).........
MAHUNKA, Sandor & Luise MAHUNKA-PAPP. Oribatids from Switzerland III
(Acari: Oribatida: Oppiidae | and Quadroppiidae). (Acarologica Genavensia
MAHUNKA, Sandor. Oribatids from Sabah (East Malaysia) VIII (Acari: Oribatida:
Dampfiellidae and Otocepheidae). (Acarologica Genavensia LXXXVI)......
MERZ, Bernhard. Two new Tephritidae (Diptera) from the Western Palaearctic
RAT BE ne Cu à eee ne LR PE PNR Mla cin 5.0.0056 0°
MUSTER, Christoph & Konrad THALER. Das Männchen von Zelotes zellensis
Grimm(Araneae:/Gnaphosidae)!.... 2... NOR fae oe ee eee
PAGES, Jean. Japygidés (Diplura) du Sud-Est asiatique n° 9. - Dicellurata Gena-
VERSO PON IN nenne release IE
POMORSKI, Romuald J. & Dariusz SKARZYNSKI. Redescription of Hymenaphorura
alticola (Bagnall, 1935) from the Alps and description of a new related
species from the Sudetes, Hymenaphorura improvisa sp. n. (Collembola:
Ouychiundae): rn Ni os ches Se ee SE EEE
PUTHZ, Volker. The genus Dianous Leach in China (Coleoptera, Staphylinidae).
2618 Contribution! to the knowledge of Steninaer .. 2.22. mer een
ROWELL, C. H. F. Review of the Lithoscirtus genus group (Orthoptera, Acrididae,
Proctolabinae)) with description of new) Species; Re CPP EC EE
Pages
81-95
165-211
3-30
335-349
107-122
233-243
283-323
277-281
871-882
601-656
721-736
49-79
675-720
883-892
579-589
737-764
657-662
419-559
793-834
INDEX DES AUTEURS
SILVA, Carla Maria Menegola da & Beatriz MOTHES. Three new species of Geodia
Lamarck, 1815 (Porifera, Demospongiae) from the bathyal depths off
Braziliankcoase SouthwestemirAtlantt Cae ee Zur OO
TANDY, Mills & Jean-Luc PERRET. The Bufo tuberosus species group with the
description of a new species from the rainforest of Cote-d’Ivoire..........
VALLAN, Denis. A new species of the genus Stumpffia (Amphibia: Anura:
Microhylidae) from a small forest remnant of the central high plateau of
Madagascar EA ee na te eer RAO wie RENE
VIT, Stanislav. Contribution à la connaissance de la famille Eucinetidae (Cole-
UE TAN) Nee Paste ERNEST meee et SEAN Mi aL EM ete a eue ve EN ON
VOGEL, Peter, Mark LAWRENCE & Ali AGHNAJ. Note sur les musaraignes
(Soricidae, Mammalia) du Parc National du Souss-Massa, Maroc.........
ZAWADZKI, Claudio H., Roberto E. Reis & Erasmo RENESTO. Allozyme discri-
mination of three species of Loricariichthys (Siluriformes: Loricariidae) from
Southe mp raza se ee Nest
XI
31-48
389-418
835-841
123-138
591-599
663-674
ICIETE SUISSE DE ZOOLOGIE
ÉUM D'HISTOIRE NATURELLE
AGMHSONIA
| RS 2000 ISSN 0035 - 418 X
SWISS JOURNAL OF ZOOLOGY
REVUE SUISSE DE ZOOLOGIE
REVUE SUISSE DE ZOOLOGIE
TOME 107 — FASCICULE |
Publication subventionnée par:
ACADEMIE SUISSE DES SCIENCES NATURELLES ASSN
VILLE DE GENEVE
SOCIETE SUISSE DE ZOOLOGIE
VOLKER MAHNERT
Directeur du Muséum d'histoire naturelle de Genève
MANUEL RUEDI
Chargé de recherche au Muséum d’histoire naturelle de Genève
CHARLES LIENHARD
Chargé de recherche au Muséum d'histoire naturelle de Geneve
Comité de lecture
Il est constitué en outre du président de la Société suisse de Zoologie, du directeur du
Muséum de Genève et de représentants des Instituts de zoologie des universités
suisses.
Les manuscrits sont soumis à des experts d’ institutions suisses ou étrangères selon le
sujet étudié.
La préférence sera donnée aux travaux concernant les domaines suivants: biogéo-
graphie, systématique, évolution, écologie, éthologie, morphologie et anatomie
comparée, physiologie.
Administration
MUSEUM D'HISTOIRE NATURELLE
1211 GENÈVE 6
Internet: http://www.ville-ge.ch/musinfo/mhng/page/rsz.htm
PRIX DE L'ABONNEMENT:
(en francs suisses)
Les demandes d'abonnement doivent être adressées
à la rédaction de la Revue suisse de Zoologie,
Muséum d'histoire naturelle, C.P. 6434, CH-1211 Genève 6, Suisse
SUISSE Fr. 225.— UNION POSTALE Fr. 230.—
ANNALES
de la
SOCIETE SUISSE DE ZOOLOGIE
et du
MUSEUM D'HISTOIRE NATURELLE
de la Ville de Geneve
tome 107
fascicule 1
2000
E à
kl GENEVE MARS 2000 ISSN 0035 - 418 X
SWISS JOURNAL OF ZOOLOGY
REVUE SUISSE DE ZOOLOGIE
REVUE SUISSE DE ZOOLOGIE
TOME 107 — FASCICULE 1
Publication subventionnée par:
ACADEMIE SUISSE DES SCIENCES NATURELLES ASSN
VILLE DE GENEVE
SOCIETE SUISSE DE ZOOLOGIE
VOLKER MAHNERT
Directeur du Muséum d'histoire naturelle de Genève
MANUEL RUEDI
Charge de recherche au Muséum d’histoire naturelle de Geneve
CHARLES LIENHARD
Chargé de recherche au Muséum d'histoire naturelle de Genève
Comité de lecture
Il est constitué en outre du président de la Société suisse de Zoologie, du directeur du
Muséum de Geneve et de représentants des Instituts de zoologie des universités
suisses.
Les manuscrits sont soumis a des experts d’institutions suisses ou étrangeres selon le
sujet étudié.
La préférence sera donnée aux travaux concernant les domaines suivants: biogéo-
graphie, systématique, évolution, écologie, éthologie, morphologie et anatomie
comparée, physiologie.
Administration
MUSÉUM D'HISTOIRE NATURELLE
1211 GENÈVE 6
Internet: http://www.ville-ge.ch/musinfo/mhng/page/rsz.htm
PRIX DE L'ABONNEMENT:
SUISSE Fr. 225.— UNION POSTALE Fr. 230.—
| (en francs suisses)
Les demandes d'abonnement doivent étre adressées
à la rédaction de la Revue suisse de Zoologie,
Muséum d'histoire naturelle, C.P. 6434, CH-1211 Genève 6, Suisse
REVUE SUISSE DE ZOOLOGIE 107 (1): 3-30; mars 2000
Description of five new species of Holoparasitus s. str. with
redescription of H. apenninorum (Berlese, 1906) and A. cultriger
(Berlese, 1906) from Italy and Spain (Acari, Gamasida, Parasitidae)
Ilinca JUVARA-BALS! & Wojciech WITALINSKI?
| Museum of Natural History, CP 6434, CH-1211 Geneva 6, Switzerland.
? Institute of Zoology, Jagiellonian University, Ingardena 6, PL-30060 Krakow,
Poland.
Description of five new species of Holoparasitus s. str. with redes-
cription of H. apenninorum (Berlese, 1906) and H. cultriger (Berlese,
1906) from Italy and Spain (Acari, Gamasida, Parasitidae). - The spe-
cimens included under the name of H. apenninorum (Berlese, 1906) in the
Berlese collection in Florence are reviewed and a lectotype for H. apen-
ninorum is designated. Five new species, H. cornutus sp. n., H. crassi-
setosus Sp. n., H. digitiformis sp. n., A. ellipticus Sp. n., H. gibber sp. n. are
described and H. cultriger (Berlese, 1906) is redescribed, all from material
from Italy and Spain. The characteristics of a new species group mallorcae
is given.
Key-words: Acari - Gamasida - Parasitidae - Holoparasitus - taxonomy.
INTRODUCTION
The gamasids belonging to the genus Holoparasitus Oudemans, 1936 are free
living, ground inhabiting predatory mites, distributed in the Holarctic region. This
genus comprises 34 species which are divided, accordingly to Juvara-Bals (1975) and
Hyatt (1987), into three subgenera: Holoparasitus s. str. (30 species), Heteroparasitus
Juvara-Bals, 1975 (3 species) and Ologamasiphis Holzmann, 1969 (1 species).
In the subgenus Holoparasitus s. str., Micherdzifiski (1969) distinguished two
species-groups: calcaratus and pollicipatus-groups, whereas the remaining species are
considered as incertae sedis. Juvara-Bals (1975) proposed another group of species,
named caesus-group. More recently Hyatt (1987) mentioned only the species-groups
recognized by Micherdzinski and designated a neotype for H. calcaratus (C.L. Koch,
1839), the type species of genus Holoparasitus.
During the last decade new taxa have been identified in the course of faunistic
and ecological programs or from acarological collections (Schmölzer, 1991, 1995a,
1995b; Vinnik, 1994; Witalinski, 1994a, 1994b: Juvara-Bals, 1995). As mentioned
above, Holoparasitus s. str. includes now 30 species. Unfortunately, some of these
Manuscript accepted 27.10.1999
4 ILINCA JUVARA-BALS & WOJCIECH WITALINSKI
species are known incompletely (one sex only) and their descriptions frequently lack
details. This rather large subgenus strongly requires a phylogenetical analysis.
However, it cannot be successfully done until the following problems are still unsolved:
- the revision of the species deposited in the collections of the Berlese, v.
Vitzthum and Willmann;
~ the re-examination of some key morphological characters omitted in early
descriptions in some species;
~ a more complete documentation about the gamasid mites in many European
regions, especially those from south-east and central Europe, is available.
The aim of this paper is to advance the knowledge of species included in the
subgenus Holoparasitus s. str. as defined in Juvara-Bals (1975). A revision of spe-
cimens labelled as A. apenninorum (Berlese, 1906) in the Berlese collection (“Berlese
Acaroteca”, Florence, Italy) showed that this is a complex of four species: one of
them we have identified as the nominal species while the three others are new species.
We also describe two additional new species from Spain and Sicily, H. gibber sp. n.
and A. ellipticus sp. n., respectively. These latter species form together with A. sicu-
lus (Berlese, 1905), H. mallorcae Juvara-Bals, 1975, H. lawrencei Hyatt, 1987 and
H. maritimus Hyatt, 1987, a well defined group of species which we named mallor-
cae-group. Additionally, we redescribe the male of A. cultriger (Berlese, 1906) based
on the single type specimen in the Berlese Acaroteca.
MATERIAL AND METHODS
The material comes from Italy and Spain. One of us (WW) was provided with a
large collection of samples from Italy collected by Prof. R. Dallai and the staff of the
Department of Evolutionary Biology, University of Siena, Italy (DEBS). Juvara-Bals
had the opportunity to identify samples of mites (family Parasitidae) from Italy
(Toscana) collected by Dr. F. Pegazanno and Dr. R. Nannelli from the Experimental
Institute of Agricultural Zoology (EIAZ), Florence, Italy. She was also working on the
genus Holoparasitus from the Athias-Henriot collection deposited in the Museum of
Natural History (MHNG), Geneva, Switzerland.
We studied material from the Berlese Acaroteca (EIAZ) which contains several
species collected generally around Florence (Toscana) and in Umbria.
Morphological terminology follows mainly Evans and Till (1979) and Van der
Hammen (1980). Setal notation for the idiosoma follows Lindquist and Evans (1965).
Measurements were made from slide-mounted material and expressed in micrometers
(um). The types are deposited in the Zoological Museum of the Jagiellonian University,
Cracow, Poland (ZMJU), in the MHNG and in the EIAZ.
Fic. |
Holoparasitus apenninorum (Berlese,1906). Male: A- chelicera, paraxial view; B-idem, antiaxial:
C-pedipalp, trochanter and femur; D-corniculus; E-leg II, femur, genu and tibia.
H. cultriger (Berlese,1906). Male: F-corniculus; G-chelicera, paraxial; H-sternogenital region and
genital lamina; I-leg II, femur, genu and tibia; J-tectum (after Berlese, 1906).
NEW SPECIES OF HOLOPARASITUS 5
6 ILINCA JUVARA-BALS & WOJCIECH WITALINSKI
SYSTEMATIC ACCOUNT
REVISION OF HOLOPARASITUS APENNINORUM (BERLESE, 1906) AND H. CULTRIGER
(BERLESE, 1906)
The material of H. apenninorum (Berlese, 1906) in the Berlese collection
named Ologamasus pollicipatus var. apenninorum consists of 28 specimens on 10
slides and is in fact a mixture of four species, 1.e.:
1. H. apenninorum (Berlese, 1906), 16 non-dissected (slide 5/19), Vallom-
brosa (Toscana, Italy), from moss; lectotype by present designation.
2. H. crassisetosus sp. n., 19 dissected (6/40), 49 2 (6/41), 688, 1%
2 deutonymphs (7/35), 26 4, 29 9 (7/37), Vallombrosa.
3. H. digitiformis sp. n., 12 (6/42), 15 (6/43), 12 (6/44), Monte Senario
(Toscana).
4. H. cornutus sp. n., 16, 12 (7/39), Bevagna (Umbria, Italy); 16, 39 2
(7/40), Monte alle Forche (Toscana).
Berlese also recognized in his material from Vallombrosa another variety,
Ologamasus pollicipatus var. cultriger. This is a valid species, Holoparasitus cultri-
ger (Berlese,1906), though only one specimen is known, a dissected male which is on
slide 5/19 together with the lectotype of H. apenninorum noted above. Some speci-
mens of Paragamasus decipiens (Berlese, 1903) and of Holoparasitus mentioned
above are mounted together: 1 2 on slide 6/44, 1 2 on slide 7/39 and 16, 19 on slide
7/40.
H. apenninorum (Berlese, 1906) Fig. 1 A-E
Gamasus (Ologamasus) pollicipatus var. apenninorum Berlese, 1906: 253 (in part).
Holoparasitus lichenis var. apenninorum (Berlese, 1887) sensu Turk 1953, Micherdzifiski
196972;
not Holoparasitus apenninorum (Berlese, 1906) sensu Juvara-Bals 1975, Acarologia, p: 400.
Type material: lectotype 1d non-dissected, slide 5/19, from moss, Vallombrosa (Tos-
cana) in Berlese Acaroteca, EIAZ; by present designation.
Our description is based on a non-dissected single male in lateral position so
that only some of the main characters could be observed.
Diagnosis. Male: excipulum absent; cheliceral movable digit with a single tooth,
fixed digit enlarged along its distal third, with 6-7 denticles; leg II with the femoral
apophysis straight and thumb-like (Fig. IE ).
Description. Male. The legs I-IV were attached only on the right side of the
animal. The others were lost but leg II could be seen in the mounting medium detached
from the animal. Gnathosoma. Movable digit of chelicera with a single tooth situated
medially. Fixed digit, enlarged along its upper third, with 6-7 denticles. Spermatotreme
in the form of a fine slit, arthrodial process setiform antiaxially and brush-like
paraxially (Fig. | A,B). Pedipalp: trochanter simple, femur with slight ventral protu-
berance located distally (Fig. 1 C). Corniculi triangular, with small protuberance on
ventral side (Fig. | D).
NEW SPECIES OF HOLOPARASITUS 7
Leg II. Main features of leg II shown in figure 1 E. Femoral apophysis straight
and thumb-like, axillary process elongated and smooth. Spur on genu and tibia conical,
located midway on ventral face. Measurements: tarsus | = 104 um; tarsus IV = 122um.
Discussion. Berlese’s (1906) original description of H. apenninorum was based
on a male and several females, from a moss sample taken at Vallombrosa. Berlese
described in detail only the male’s chelicera but presented figures of three different
chelicerae as well as of the epistome, and of the male’s leg II. The drawing of the
chelicera shown in the fig. 15a, tab. XIII. (6 from Vallombrosa) corresponds to the
original description which specified that “digitus mobilis basi latiusculus, externe
gibbosus, dente magno, unico...”; the characteristics of this type of chelicera can be
seen only on the non-dissected male of the slide 5/19. Berlese attributed to H. apenni-
norum two other chelicerae (Fig. 15, Fig. 16, tav. XIII) which have two teeth on the
inner margin of the digitus mobilis. He also mentioned a figure 16a which in fact does
not exist. Micherdzinski (1969) paid also attention to the similarity between Berlese’s
initial description and only one of the chelicerae illustrated. Berlese’s figure of the
male’s leg II corresponds either to the leg of H. digitiformis or to that of H. crassiseto-
sus, two species found in the same area as H. apenninorum. The triangular epistome
figured by Berlese can not be seen any more because of the lateral position of the
animal.
The characteristics of the female endogynium was neither described nor drawn
in detail; figure 7, tab. XIX refers either to H. digitiformis or to H. crassisetosus which
have very similar females. Berlese’s material also contained H. crassisetosus from
Vallombrosa and H. digitiformis from Monte Senario. Unfortunately, we did not find
another male of H. apenninorum or a corresponding female. However, it is obvious that
the male mounted on slide 5/19 is a syntype. We designate it as the lectotype of H.
apenninorum (Berlese, 1906).
Specimens identified by Juvara-Bals (1975) as H. apenninorum belongs in fact
to H. cornutus sp. n. The species was misidentified because at that time it was not
recognized that original description of Berlese related to a mixture of species.
H. cultriger (Berlese, 1906) Fig. I F-J
Type material: 16 holotype, dissected (slide 5/19), Vallombrosa (Toscana, Italy), from
moss, Berlese Acaroteca (EIAZ).
Diagnosis. Male: excipulum present; cheliceral movable digit with two little
denticles, fixed digit markedly longer than movable one, blade-like and toothless; leg II
with tibia bearing two processes: dorsally one rounded located near proximal margin
and ventally an elongated situated near distal margin (Fig. 1 D.
Description. Male. The length of the idiosoma could not be measured because
the single specimen is in pieces. The few characteristics observed are shown in figure 1.
Sternogenital region reticulated, with slightly sclerotized excipulum; anterior margin of
genital lamina with several small denticles in the middle (Fig. | H).
Gnathosoma. Tectum trispinate, central prong well developed (Fig. 1 J). Corni-
culi triangular, each with ventral protuberance (Fig. 1 F). Chelicera as in figure 1 G:
8 ILINCA JUVARA-BALS & WOJCIECH WITALINSKI
fixed digit blade-like and toothless, markedly longer than movable one; movable digit
with two little denticles subapically.
Legs. Armature of legs II as in figure 1 I. Femoral apophysis strong, thumb-like
and curved; axillary process short and rounded. Genual process large, plump or
rounded, situated in distal half on ventral face. Tibia with two protuberances: regular
process elongated and located ventrally close to distal margin of segment, an extra
process larger, located dorsally on proximal third of segment. Ventral face of trochanter
IV with protuberance. Measurements: tarsus I = 138 um, tarsus IV = 150 um.
Discussion. Berlese described this species on the basis of a single male in poor
condition. The type specimen is together with that of H. apenninorum on the same
slide. The two types may be easily distinguished: H. apenninorum is in lateral view and
non-dissected while H. cultriger is in several pieces. The most valuable character
separating H. cultriger from the other species with an excipulum is the special blade-
shaped fixed digit of the chelicera.
Holoparasitus crassisetosus sp. n. Figs. 2,3
Type material: 13 holotype, 486 d, 669 © paratypes, Vallombrosa (Toscana, Italy)
(alt. 960 m), coniferous forest (Abies alba), 6.11.1982. Collected by the staff of DEBS. The
holotype as well as 46 d and 59 © paratypes have been deposited in MHNG, 446 4 and
612 2 have been deposited in ZMJU.
Other material examined: Berlese Acaroteca (EIAZ), 19 (slide 6/40), 49 2 (6/41),
288,222 (7/37), 584,299, 2 deutonymphs (7/35). All this material was collected in
Vallombrosa, but the habitat was not specified.
Diagnosis. The species is recognisable by very thick seta pv! on tibia IV in both
sexes (Fig. 2 A,B).
Description. Male. Idiosoma brown, well sclerotized. Dimensions of idiosoma:
555-585 x 365-380 um; L/W (length/width) factor 1.46-1.56, N=10. Podonotal region
with 20 pairs of setae, j1 = 38-40 um, other 19-22 um; opisthonotal region with 30
pairs of shorter setae, 10-13 um.
Ventral side (Fig. 2 C). Genital lamina large, located in a well sclerotized
concavity, its lateral sides forming triangular sharp projections, its anterior edge with
median pleated membrane and two lobes; postero-lateral edges of genital lamina with
two well sclerotized protrusions.
Male genital orifice, flanked by triangular presternal shields and provided with
large rectangular microsclerite bearing tritosternum. Reticulation of sternogenital
region regular; between anterior margin and close to sternal setae 1 (stl), sternal shield
more heavily sclerotized and its reticulation slightly convex; length of sternal setae
about 42 um. Opisthogastric region with 8 pairs of setae (26-39 um).
Gnathosoma. Tectum trispinate, with long, broad central prong (Fig. 3 I). Corni-
culi with paraxial margin forming rounded lamellar protrusion (Fig. 2 D,E). Hypo-
gnathal groove with 11 complete rows of denticles; palpcoxal setae pilose, hypo-
stomatic setae simple (Fig. 2 D). Chelicera (Fig. 2 F,G). Both digits short and robust.
Fixed digit toothless, with pilus dentilis flanked by two convex laminae. Movable digit
with two teeth. Arthrodial membrane at the base of movable digit with well developed
NEW SPECIES OF HOLOPARASITUS 9
Fic. 2
Holoparasitus crassisetosus sp. n. Male: A-tibia IV; C-sternogenital region and genital lamina;
D-gnathosoma, palptrochanter and palpfemur; ventral; E-corniculus; F-chelicera, antiaxial:
G-idem, paraxial; H-leg II, femur, genu and tibia, antiaxial; I-tibia, ventral; J-basis tarsus and
cuticular edge of tibia; Female: B-tibia IV.
10 ILINCA JUVARA-BALS & WOJCIECH WITALINSKI
brush-like process on paraxial side and a lesser developed one on antiaxial side,
synarthrodial membrane short and wedge-shaped.
Pedipalps. Trochanter with proximal seta (v1) simple and distal seta (v2) pilose.
Femur with tubercle located on ventral side close to anterolateral seta. Anterolateral
seta of femur spatulate and pectinate on inner side (Fig. 2 D); anterolateral setae of
genu spatulate.
Legs. Coxae II with ridge of 5 denticles (Fig. 3 J). Leg II armed as shown in
figure 2 H-I. Femoral apophysis finger-shaped and straight, axillary process curved
towards the femur. Spur on genu small, round, located ventrally in the middle.
Apophysis on tibia low and long, with slightly convex margin, attached ventrally and
reaching with its rounded distal end the margin of segment; the cuticle of anterodistal
edge of tibia with several fine furrows. Trochanter IV with flattened protuberance
situated medioposteriorly (Fig. 2 K). On tibia IV seta pv! conspicuous, very thick with
a dentate end (Fig. 2 A). Measurements: tarsus I = 139-144 um; tarsus IV = 146um.
Female. Idiosoma brown, well sclerotized. Dimensions of idiosoma: 630-665 x
445-475um (L/W factor 1.36-1.45, N=10). Podonotal setae: j1 = 39 um, other setae 19-
26um; opisthonotal setae 12-18 um.
Ventral side (Fig. 3 A). Fused presternal shields forming smooth ribbon,
partially connected to lateral platelets. Anterior margin of sternal plate sometimes with
incisions of a soft cuticle, laterally to setae stl. Reticulation of sternal shield with two
prominent lines delimiting a slightly more sclerotized anterior region of sternum: one
line V-shaped extending medially and second line running close to second pair of pores
(pst2). Margin of exopodal plate facing coxa I sclerotized.
Genital region. Epigynial plate heptagonal, anterior margin with a middle
triangular apex and two lateral spines curved antiaxially; subapical structure small and
oval, with lateral rounded protrusions extending beyond epigynial margin (Fig. 3 C).
Paragynia rectangular with elliptical thickenings in front of coxae IV and not extending
beyond paragynium edge, paragynial posterior protrusions elongated (Fig. 3 B).
Endogynium oval, 22 x 29 to 24 x 31 um. Distance between endogynial margin and
coxa IV nearly two times larger than endogynium length (56-59 um). Shape of
endogynium cup-like, with anterior margin protruding into dorsally directed appendage,
which looks like hollow tube (Fig. 3 D). Gland pores behind coxa IV large. Opistho-
gastric region with 8 pairs of setae, their length 36-39 um.
Gnathosoma. Tectum trispinate, central prong long and narrow, lateral prongs
small; minute additional spines discernible between central and lateral prongs (Fig.
3 E). Corniculi conical. Hypognathal groove with 6-8 rows of denticles ending at
palpcoxal setae level; some lateral lines present, starting from hypognathal groove;
cuticular ridges between palpcoxal setae serrated (Fig. 3 F). Palpcoxal setae slightly
pilose, hypostomatic setae simple.
Chelicera. Fixed digit with two teeth in front of pilus dentilis, a third, smaller
tooth frequently present between them; two teeth and a lamella with slightly concave
edge located behind pilus dentilis. Movable digit with three teeth (Fig. 3 G).
Pedipalps. Proximal seta of trochanter (v1) simple, distal seta (v2) pilose (Fig. 3
F), anterolateral seta of femur spatulate and pectinate on one side, and anterolateral
setae of genu spatulate. Femur with poorly visible tubercle located anterolaterally.
NEW SPECIES OF HOLOPARASITUS 11
FIG. 3
Holoparasitus crassisetosus sp. n. Female: A-presternal and sternal shields; B-paragynium; C-
epigynium; D-endogynium; E-tectum; F-gnathosoma and palptrochanter, ventral; G-chelicera,
paraxial; H-trochanter IV, ventral. Male: I-tectum; J-coxa II, denticulated ridge.
12 ILINCA JUVARA-BALS & WOJCIECH WITALINSKI
Legs. Coxa II with a ridge of about 5 denticles on anterior face. Tibia IV with
very thick pv1 seta, its end dentate (Fig. 2 B). Trochanter IV with rounded protuberance
located on proximal third of posterior face (Fig. 3 H).
Measurements: tarsus I = 144-154 um; tarsus IV = 156-163 um.
Discussion. Males and females of H. crassisetosus are very similar to those of
H. digitiformis. From other Holoparasitus species known to date these species differ in
the following features. In males, excipulum is absent, tectum trispinate, movable and
fixed digits of chelicera with 2 and 0 teeth, respectively. Corniculi posses a lamellar
dilatation or indentation on the paraxial margin. In females, presternal plates and lateral
platelets accreted, the endogynium soup-spoon-shaped. The most easily recognized
feature which distinguishes H. crassisetosus and H. digitiformis is the presence of the
transformed pvl seta of tibia IV in both sexes of the former species. A detailed
discussion of the most important differences between the two species is given at the end
of the description of H. digitiformis.
Holoparasitus digitiformis sp. n. Figs 4, 5
Type material: 15 holotype, 526 8, 392 © paratypes, from litter of oak forest (Quercus
cerris, Q. pubescens) and 68 6d and 729 2 paratypes, from litter of pine forest (Pinus nigra,
Erica arborea, Juniperus communis). The two forests are situated at Selvapiano (Commune di
Rufina, Toscana), 25 km away from Florence at an altitude of 200 m. The two habitats are about
300 m apart and are situated on the same side of the mountain, facing south-west. The material
was collected from February 1971 to April 1974 by Dr. R. Nannelli and Dr. F. Pegazzano
(EIAZ).
Other material examined: 193 4, 159 ?, Catena del Marghine (Marghine Range),
Sardinia, from moss and lichens in yew (Taxus baccata) and oak forests along the road to
Bolotana, alt. ca 1000 m, 30.03.1977, collected by the staff of the DEBS and deposited in ZMJU.
12 (slide 6/42), 18 (6/43), 12 (6/44), Monte Senario, Toscana, Berlese Acaroteca
(EIAZ). The material has been deposited as follows: 1d holotype, 326 d, 199 © paratypes from
oak forest and 48d d, 529 © paratypes from pine forest — in MHNG; 203 d, 209 © paratypes
from both habitats - in ZMJU; 203 4,209 © paratypes from both habitats - in EIAZ.
Diagnosis. In both sexes all setae on tibia IV of normal appearance. Male:
excipulum absent; movable digit of chelicera with two denticles, fixed digit edentate,
slender, finger-like, with slightly dilated apex. Female: epigynium with subapical small
ovoid microsclerite and hyaline wing-like protrusions extending anterolaterally (Fig. 5
D). Endogynium small, oval, inverted cup-like, with anterior margin protruding into a
cuticular solid appendage directed more or less dorsally (Fig. 5 B,C,E).
Description. Male. Idiosoma well sclerotized. Dimensions of idiosoma: speci-
mens from Selvapiano, Toscana, oak forest: length 576-617 um; idem, specimens from
pine wood, length 634-641 um. Specimens from Sardinia, idiosoma size 585-635 x
405-430 um (L/W factor 1.45-1.57, N=10). Podonotal region with 21 pairs of seta,
j1=39 um, the others 26-28 um opisthonotal region with about 30 pairs, 19-24 um.
Ventral side. Genital lamina large, situated in well defined concavity of heavily
sclerotized anterior margin of sternal shield; anterior margin of the lamina with two fine
lobes separated by a concavity and two lateral triangular prongs. Between lateral prongs
and anterior lobes, on their inner face, two well sclerotized prominences. Behind genital
lamina a rectangular microsclerite with a rounded lobe ventrally supporting the base of
NEW SPECIES OF HOLOPARASITUS
Fic. 4
Holoparasitus digitiformis sp. n. Male: A-genital lamina; B-sternogenital shield; C-tectum;
gnathosoma and palptrochanter, palpfemur, ventral; E-chelicera; F-idem, paraxial; G-leg
femur, genu and tibia, paraxial; H-femoral apophysis, antiaxial; I-trochanter IV, ventral.
ID:
II,
14 ILINCA JUVARA-BALS & WOJCIECH WITALINSKI
tritosternum (Fig. 4 A). Sternogenital shield with scale-like reticulation and with
marked line (inverted V) behind second pair of sternal setae; length of sternal setae 39-
46 um (Fig. 4 B). Large gland pore behind coxae IV. On opisthogaster 8 pairs of setae,
24-36 um.
Gnathosoma. Tectum trispinate, lateral prongs small, the central one large, broad
at the base and pointed (Fig. 4 C). Corniculi with small prominence situated paraxially;
hypognathal groove with 10-11 rows of denticles; palpcoxal setae finely pilose, hypo-
stomatic setae simple (Fig. 4 D).
Chelicera (Fig. 4 E,F). Fixed digit edentate, slender, finger-like; its apex slightly
dilated and sometimes curved. Movable digit with two denticles, the distal one
sometimes appearing much more paraxially, synarthrodial membrane rounded. Sperma-
todactyl large, with a median concavity; arthrodial membrane with short brushy process
paraxially and setiform margin antiaxially.
Pedipalps. Palptrochanter with seta vl thin and simple, whereas v2 thicker and
pilose. Palpfemur with anterolateral seta pectinate on one edge; palpgenual anterolateral
setae simply spatulate. Palpfemoral segment swollen distally, with tubercle located on
ventral side, close to al seta (Fig. 4 D).
Legs. Coxa I with a ridge bearing 6-7 denticles, situated paraxially close to
distal margin. Coxa II with ridge of 8-10 denticles (Fig. 5 J). Armature of leg II illus-
trated in figure 4 G.
Femoral apophysis relatively long, finger-shaped, with curved apex and a small
tubercle on its base (Fig. 4 H); axillary process curved. Genu with small rounded spur
situated medioventrally. Tibia with two spurs: one low and elongated, reaching distal
margin, another smaller, located anterolaterally and extending beyond segment margin.
Posterior face of trochanter IV with a small tubercle under seta pv (Fig. 4 I).
Measurements: specimens from Toscana: tarsus I = 156-158 um; tarsus IV =
168-173 um.
Female. Idiosoma well sclerotized, brown coloured. Dimensions of idiosoma:
660-710 x 490-530 um (L/W factor 1.28-1.42, N=10). Podonotal region with 21 pairs
of setae, }1=36um, the others 16-20 um; opisthonotal region with 33 pairs of short setae
(13-16 um).
Ventral side. Accreted presternal shields forming a smooth ribbon-shaped plate,
more narrow medially; lateral presternal platelets usually free. Anterior margin of
sternal plate with two less sclerotized concavities, frequently flanking bases of setae
stl; reticulation of sternum easily discernible with prominent arched transverse line
passing close to pores pst2 (Fig. 5 A); length of sternal setae from 46 um (stl) to 65 um
(st3). Anterior edge of exopodal shield facing leg I thickened.
Genital region. Paragynial shield with sclerotized elliptical thickening in front of
coxa IV (Fig. 5 B.C). Postero-lateral protrusions long and narrow, ending with rounded
or sometimes pointed edge. Metagynial sclerites with paraxial margins convex.
Epigynial shield with anterior margin formed by a triangular central apex and two large
lateral prongs curved antiaxially; subapical epigynial structure with small ovoid
microsclerite and hyaline wing-like protrusions extending anterolaterally (Fig. 5 D).
Endogynium (Fig. 5 B,C,E) small, oval (20 x 27 um to 27 x 33 um), inverted cup-like,
NEW SPECIES OF HOLOPARASITUS 15
li 0,1mm. A,B,C,D,J
SU Ma
me “i 005mm. EI
Fic. 5
Holoparasitus digitiformis sp. n. Female.: A-presternal and sternal shields; B-paragynia and
endogynium, dorsal; C-idem, ventral; D-epigynium; E-endogynium; F-tectum; G-hypognathal
groove and corniculus; H-chelicera, antiaxial; I-coxa II, denticulated ridge. Male: J-coxa II,
paraxial.
16 ILINCA JUVARA-BALS & WOJCIECH WITALINSKI
with anterior margin protruding into a cuticular solid appendage directed more or less
dorsally; poorly visible scale-like fine hyaline flap covering endogynium on ventral
side. Gland pores behind coxa IV large. Opisthogastric region with 8 pairs of setae,
their length about 39 um.
Gnathosoma. Tectum usually trispinate, with central prong sharply pointed,
sometimes an extra spine present on one side (Fig. 5 F). Corniculi conical. Hypognathal
groove with 8-11 rows of denticles, those located posterior to the level of palpcoxal
setae frequently reduced or absent, some lateral lines present between hypostomatics
and palpcoxal setae (Fig. 5 G ). Palpcoxal setae pilose, hypostomatics simple.
Chelicera. Fixed digit with two teeth in front of pilus dentilis, two more teeth
and a thin, concave cuticular edge behind pilus dentilis. Movable digit with three teeth
(Fig. 5 H).
Pedipalps. Trochanter ventral setae finely pilose; anterolateral seta of femur
spatulate and pectinate on one edge; anterolateral setae of genu simply spatulate. Palp-
femur swollen distally, with tubercle located on ventral side close to anterolateral seta.
Legs. Leg structure and setation unremarkable. Coxa I with row of about 10
little denticles on distal paraxial margin. Coxa II with ridge of 8 to 10 fine denticles
situated on anterior face (Fig. 5 I). Measurements: tarsus I = 154-163 um; tarsus IV =
168-175 um.
Discussion. H. digitiformis is similar to H. crassisetosus in respect to the general
structure of the genital region in the female and to the shape of the armature of leg II in
the male.
The endogynium in both species is small and inverted cup-shaped, with the
anterior margin forming a cuticular protrusion directed dorsally, thus similar in
appearance to a soup-spoon. In H. crassisetosus, this protrusion is relatively shorter (as
compared with the size of endogynium) than in H. digitiformis and appears as a hollow
tube. The metagynial sclerites are oriented roughly parallel in both species, but in A.
digitiformis their lateral margins are convex, whereas in H. crassisetosus they are rather
concave. The subapical epigynial structure has prominent wing-like protrusions in H.
digitiformis but only small round protrusions in H. crassisetosus .
Males in both species have a very similar armature of leg II, especially on the
femur and genu: on the tibia, however, in A. digitiformis the additional prominent
anterolateral tubercle is present, whereas in H. crassisetosus the cuticle of corres-
ponding place shows fine furrows only. A very pronounced difference can be seen in
the shape of the male corniculi: H. digitiformis has a small protuberance on the paraxial
margin, whereas in H. crassisetosus this margin extends prominently forming a thin
rounded lamella.
In both sexes, the most important character for differentiation of these two
species remains the form of seta pv! of tibia IV: it is narrow and simple in H. digi-
tiformis but thick and notched at the end in A. crassisetosus. H. digitiformis was found
in the surroundings of Florence (Toscana) in coniferous forest as well as in Sardinia
where it occurs in soil of yew and oak forests. H. crassisetosus was collected, till now,
only from Vallombrosa (Toscana), where it lives in coniferous forests.
NEW SPECIES OF HOLOPARASITUS 17
TAR
0,05mm.
Fic. 6
Holoparasitus cornutus sp.n. Male: A-scutum sternogenital and genital lamina; B-tectum:
C-tectum (Bevagna-Umbria, Berlese Acaroteca); D-gnathosoma and palptrochanter, ventral;
E-chelicera, antiaxial; F-leg II, femur, genu and tibia; G-tibia II, ventral (Bevagna-Umbria,
Berlese Acaroteca); H-coxa II, denticulate ridge; I-trochanter IV, ventral; J-idem, dorsal.
18 ILINCA JUVARA-BALS & WOJCIECH WITALINSKI
Holoparasitus cornutus sp. n. Figs. 6, 7
H. apenninorum sensu Juvara-Bals 1975: 400: nec Berlese, 1906.
Type material: 13 holotype, 26 4, 42 2 paratypes, Vallombrosa (Toscana, Italy), litter
in beach forest, 19.09.1971: 148 &, 119 2 paratypes, Vallombrosa, moss in coniferous forest
(Abies alba), alt. 960 m, 6.11.1982. All paratypes collected by the staff of DEBS. The holotype
and 14,29 2, 2 deutonymphs have been deposited in MHNG, whereas 166 6, 132 © in ZMJU.
Other material examined. Italy: 18, 12, Monte Amiata near Siena, Toscana, litter in
beach forest, alt. ca. 900 m; 49 9, Monte Amiata, litter in chestnut forest, alt. ca. 800 m,
17.09.1990, leg. W. Witalinski, deposited in ZMJU. 16, 32 © (slide 7/40), Monte alle Forche,
Toscana, and 15,1% (slide 7/39), Bevagna, Umbria, Berlese Acaroteca (EIAZ).
Romania: 16, 19, Cerbului Valley, Busteni, Bucegi Mountain, Meridional Carpathians,
sawdust and litter, beech and spruce forest, 5.6.1967, leg. I. Juvara-Bals, deposited in MHNG.
Diagnosis. The species can be recognized by a lamellar process ending with one
or two spurs, situated more or less dorsally at the distal margin of trochanter IV in both
sexes. Male: excipulum absent; two incisions in sclerotized cuticle behind each group
of hypostomatic setae; folded cuticle at the level of legs IV. Female: endogynial sack
rounded, with two large, connected teeth on each side .
Description. Male. Idiosoma well sclerotized, its dimensions (Italy) 672-715 x
452-510 um (L/W factor 1.42-1.48, N=5). Podonotal setae: j1=38 um, other setae 20-
26 um; opisthonotal setae shorter, 13-19 um.
Ventral side (Fig. 6 A). Genital lamina with serrated anterior margin. Sternal
shield reticulated with a slight median prominence distally; two ridges run from this
prominence posteriorly to sternal setae st2 towards the margin of the shield. At the level
of coxa IV, cuticle folded in a strongly sclerotized line. Length of sternal setae from 48
um (stl) to 40 um (st3). On opisthogastric region 8 pairs of setae, their length from 42
um (Jv1) to 26 um (Jv5).
Gnathosoma. Tectum trispinate with central prong triangular, long, and in the
specimen from Bevagna, with a little denticle on its tip; lateral prongs tiny (Fig. 6 B,C).
Hypognathal groove provided with 11 well dentated rows; gnathosomal setae pilose,
except simple hypostomatic seta 3. Behind hypostomatic setae there are incisions in
sclerotized cuticle (Fig. 6 D). Corniculi conical. Chelicera (Fig. 6 E). Fixed digit eden-
tate and with truncate apex; movable digit with 2-4 small subapical teeth. Arthrodial
membrane formed by a short brush-like processes paraxially and a setiform fringe
antiaxially.
Pedipalps. Palptrochanter with seta vl simple and v2 pilose; palpfemur with
distal protuberance on its ventral face.
Legs. Coxa II with ridge bearing 9 denticles located on anterior face (Fig. 6 H).
Armature of leg II shown in figure 6 F. Main femoral apophysis long, finger-like, its
axillary spur rounded. Spur on genu rounded. Tibia with median saddle-like spur and
little distal protuberance situated anterolaterally (Fig. 6 F,G). Dorsal or posterodorsal
face of trochanter IV with distal lamellar process ending with one or two spurs;
proximal posterolateral face with a protuberance (Fig. 6 I,J). Gland pores behind coxae
IV located in a normal cuticle. Measurements: tarsus I = 143 um (specimen from
Romania) and 173-177 um (specimens from Italy); tarsus IV = 173 um (Romania) and
184-193 um (Italy).
NEW SPECIES OF HOLOPARASITUS 19
0,05mm.
le]
FIG.7
Holoparasitus cornutus sp. n. Female: A-presternal and sternal shields; B-paragynium; C-
epigynium; D-endogynium (2 from Rumania); E-idem, (9 from Italy); F-tectum; G-hypognathal
groove and corniculi; H-palptrochanter and palpfemur, antiaxial; I-chelicera, paraxial; J-tro-
chanter IV, ventral; K-coxa II, denticulate ridge; L-gland gv2.
20 ILINCA JUVARA-BALS & WOJCIECH WITALINSKI
Female. Dimensions of idiosoma: 580 x 430 um (L/W factor 1.35), specimen
from Romania; 742-806 x 550-580 um (L/W factor 1.34-1.41, N=5), specimens from
Italy. Podonotal setae from 13 um to 39 um, j1 = 36-38 um; opisthonotal setae shorter,
about 13 um. ;
Ventral side. Presternal shield smooth, ribbon-like, with two lateral excres-
cences, sternal shield with transverse ridge at level of pst2, some specimens with two
lightly sclerotized indentations in front of setae stl (Fig. 7 A). Sternal setae slender,
55-60 um. Paragynial shield provided on its dorsal face with an elliptical thickening in
front of coxa IV and on its inner ventral margin with one or two denticles; metagynial
sclerite elongated, paragynial posterior protrusion elongated and rounded, paragynial
lobe trapezoidal (Fig. 7 B). Epigynium heptagonal; its apex transparent, its subapical
structure formed by a fine rounded margin and a sclerotized three-lobed thickening
(Fig. 7 C). Endogynium: a rounded sack with two large, connected teeth on each side,
covered by a scale flap ventrally, sometimes provided with denticles on inner walls;
dorsally, its cuticle with many pores arranged in ca. 8 rows of muscle-prints (Fig. 7
D,E.). Gland pores behind each coxa IV located in a normal cuticle (Fig. 7 L). On
opisthogaster 8 pairs of setae, their length from 40 um (Jv1) to 24 um (Jv5).
Gnathosoma. Tectum trispinate, lateral prongs tiny (Fig. 7 F). Hypognathal
groove with 9-10 lines, anterior dentate, posterior bare; external posterior hypostomatic
seta simple, other gnathosomal setae slightly pilose. Corniculi conical (Fig. 7 G).
Chelicera. Fixed digit with 3 minute denticles in front of pilus dentilis and
behind it two teeth and a cuticular edge (Fig. 7 I).
Pedipalps. Trochanter with seta vl simple and seta v2 slightly pilose. Femur
with small protuberance near al seta (Fig. 7 H).
Legs. Coxa IT with ridge bearing about 6 denticles (Fig. 7 K). Trochanter IV
provided proximally with rounded protuberance on posterolateral face and with lamel-
lar process, pointed in one or two spurs, dorsally or posterodorsally on distal margin of
segment (Fig. 7 J). Measurements: tarsus I = 145 um (specimen from Romania) and
164-173 um (specimens from Italy); tarsus IV = 166 um (Romania) and 189-196 um
(Italy).
Discussion. Juvara-Bals (1975) identified and described under the name of
H. apenninorum (Berlese) specimens of Holoparasitus collected in the surroundings of
Siena (Italy) and in Bucegi Mountain, Meridional Carpathians (Romania). She thought
that the material belonged to H. apenninorum according to characteristics of male
chelicera. In fact, Berlese (1906) had drawn two types of chelicera for O. pollicipatus
var. apenninorum, as already discussed in the comments to the redescription of that
species. Figure 16, tab. XIII by Berlese (1906) was a drawing of a specimen from
Monte Senario which is the same as H. cornutus, the new species described above. The
specimens of A. cornutus from Romania and Italy do not exhibit significant differences
(Juvara-Bals, 1975). The male genital lamina is not serrated in specimens from Roma-
nia but instead it has only two central denticles. The number of teeth on the movable
digit is 2 in Romanian versus 3 in Italian specimens and the shape of the tectum is
slightly variable. H. cornutus is characterised especially by the lamellar process situated
more or less dorsally on margin of trochanter IV in both sexes. Other characteristics
NEW SPECIES OF HOLOPARASITUS 2]
FIG. 8
Holoparasitus gibber sp. n. Male: A-chelicera, paraxial; B-idem, antiaxial; C-base of movable
digit; D-tectum; E-gnathosoma and palptrochanter, ventral; F-corniculus; G-palptrochanter and
palpfemur; H-genital lamina; I-coxa I, paraxial; J-coxa II, denticulate ridge; K-leg II, femur, genu
and tibia.
22 ILINCA JUVARA-BALS & WOJCIECH WITALINSKI
such as the shape of the chelicera and armature of leg H in the male, as well as the
particularities of the endo- and epigynium of the female distinguish this species from
the other species belonging to the complex O. pollicipatus var. apenninorum sensu
Berlese 1906. We also have to mention the species H. hemisphaericus (Vitzthum, 1923)
described on the basis of a single female found in Austria. This species was considered
by Juvara-Bals (1975) to be synonymous with H. apenninorum sensu Juvara-Bals,
1975. H. cornutus was compared with the type of H. hemisphaericus kindly sent to the
senior author by Dr. L. Tiefenbacher (Munich, Germany). H. hemisphaericus is a valid
species which differs from H. cornutus by the shape and size of the dorsal protuberance
located distally on trochanter IV, by the absence of the proximal protuberance on
trochanter IV, as well as by the paragynia, characterised by thickenings at paraxial
margins but devoid of any thickening in front of coxa IV. Unfortunately, this species is
known only on the basis of one female and on the single specimen available the
characteristics of the epigynium and those of the endogynium cannot be properly
observed.
Holoparasitus gibber sp. n. Figs 8, 9
The species is described from permanent slides of the Athias-Henriot collection. The
length of the idiosoma cannot be measured because the specimens are crushed.
Type material: 13 holotype, 26 4, 32 2, 1 deutonymph, paratypes, Los Palacios (near
Seville, Spain), sifting of flood litter, 22.02.1951, leg. H. Franz, and 48 8,4? 2, 1 deutonymph
paratypes, surroundings of Carmona (near Seville, Spain), in the vicinity of Finca Alamaja, black
soil from decayed plants, 22.02.1951, leg. H. Franz. All types in collection of Athias-Henriot
deposited in MHNG.
Diagnosis. Male: excipulum absent; mobile digit of chelicera with | tooth, fixed
digit with subapically located 6 denticles plus one (Fig. 8 A,B); hypostome extended
between corniculi. Female: presternal plate with anterior margin serrated; endogynium
circular in outline with two prolongations of posterior margin reaching anterior edge of
endogynium, lateral sides of endogynial sack with 1-2 denticles (Fig. 9 D).
Description. Male. Idiosoma. Cuticle yellow-brownish, moderately sclerotized.
Setae on holodorsal shield from 36 um (j1) to 13-15 um in opisthonotal region.
Ventral side. Sternal shield reticulated, without a particular pattern; length of
sternal setae 24 um (st2) to about 36 um (stl, st4). Genital lamina with large central
rounded process (Fig. 8 H). Opisthogaster with 8-9 pairs of setae, their lengths about
20 um. Simple gland pore behind coxae IV.
Gnathosoma. Tectum with three prongs, the median much larger and broader
than lateral ones (Fig. 8 D). Hypostome extended between corniculi; hypognathal
groove provided with 9 rows of very fine denticles: hypostomatic and palpcoxal setae
simple. Sclerotized cuticle with incisions behind hypostomatics; internal malae covered
by a trapezoidal extension (Fig. $ E). Corniculi slender and conical (Fig. 8 F).
Chelicera (Fig. 8 A-C). Fixed digit straight and narrow, its apex slightly curved;
along its upper third, internal paraxial edge provided with 6 small denticles and
antiaxial edge only with one big tooth; pilus dentilis located between these two edges.
Movable digit with big tooth medially on internal margin and a characteristic rounded
NEW SPECIES OF HOLOPARASITUS
(NO)
Ww
hump on external (ventral) margin. Spermatotreme slender, ending distally at the level
of tooth: the basal part of the digit with well sclerotized enlargement and a little
denticle. Arthrodial membrane developed into brush-like process.
Pedipalps. Trochanter with protuberance located between slightly pilose setae v1
and thick and pilose v2; femur provided with small rounded protuberance, seta al
slightly spatulate (Fig. 8 G).
Legs. Distal margin of coxa I dentate, coxa II with ridge bearing 7-8 denticles
and an extra basal denticle (Fig. 8 I,J). Spurs on leg II as in figure 8 K: femoral
apophysis short and thumb-like and axillary process trapezoidal; triangular spur located
distally on genual margin; tibial apophysis elongated, situated medially on anterolateral
face. Measurements: tarsus I = 159-169 um; tarsus IV = 164-169 um.
Female. Idiosoma. Colour yellow-brownish; setae on podonotal region from 46
um (j1) to 26 um; setae on opisthonotal region shorter, ca. 13 um.
Ventral side. Presternal plate entire, ribbon-like, with the anterior margin
serrated; lateral presternal platelets free. Sternal shield reticulated with a longitudinal
granular strip medially (Fig. 9 A), length of sternal setae about 52 um. Paragynial
shields reticulated, metagynial sclerites oval (Fig. 9 B). Epigynium heptagonal, its
anterior margin with triangular median apex and two lateral prominent spines:
subapical epigynial structure formed by a weakly sclerotized rectangle and a hyaline
structure stretching beyond epigynial margin in the form of two little wings (Fig. 9 C).
Endogynium cup-shaped and circular in outline. Its posterior margin protrudes forward
to form two long, sinuous prolongations with rounded tips reaching anterior edge of
endogynial sack; lateral sides with 1-2 denticles (Fig. 9 D). Opisthogastric shield with 9
pairs of setae, their length from 42 um (Zv1) to 24 um (Jv5).
Gnathosoma. Tectum trispinate with long median prong and two tiny lateral
prongs; dorsal face slightly granular (Fig. 9 E). Hypognathal groove with 9-10 rows of
denticles, last four oligodentate; palpcoxal seta pilose, anterior hypostomatic seta
slightly barbed, posterior setae simple (Fig. 9 F).
Pedipalps (Fig. 9 G). Border of trochanter thickened between pilose setae vl and
v2; distal margin of femur with small tubercle. Chelicera (Fig. 9 H) similar to that of H.
ellipticus: fixed digit with five teeth and a long, thin cuticular ridge, movable digit with
three teeth.
Legs. Structure and setation unremarkable. Denticulate ridge on coxa II as in the
male (Fig. 9 I). Measurements: tarsus I = 156-163 um; tarsus IV = 173-185 um.
Discussion. Adults of H. gibber are similar to those of H. mallorcae and of
H. ellipticus. All these species have in common: denticulation of the anterior margin of
the female’s presternal shield, the shape of the endogynium and the pattern of male’s
second leg armature; but they differ mainly in the shape of the male’s chelicera and
tectum as well as in the ferm of the female’s epigynium. See comments following the
description of H. ellipticus.
Holoparasitus ellipticus sp. n. Fos OM
Type material: 15 holotype, 116 8,9% © paratypes, Milazzo, Sicily, Italy from litter of
olive trees, 24.03.1972; 18, 29 2 paratypes, Monti Peloritani, Sicily, from humus and litter in
24 ILINCA JUVARA-BALS & WOJCIECH WITALINSKI
Fic. 9
Holoparasitus gibber sp. n. Female: A-presternal and sternal shields; B-paragynium; C-epigy-
nium; D-endogynium; E-tectum; F-hypognathal groove and corniculus; G-palptrochanter and
palpfemur: H-chelicera, paraxial; I-coxa II, denticulate ridge.
oak forest, 26.03.1972. The material was collected by the staff of the DEBS. The holotype and
36 d, 69 9 have been deposited in MHNG, whereas 98 8,52 2 in ZMJU.
Diagnosis. Male: excipulum absent; mobile and fixed digits of chelicera with
many denticles (Fig. 10 D,E); hypostome evidently extended between corniculi.
NEW SPECIES OF HOLOPARASITUS DS
Fic. 10
Holoparasitus ellipticus sp. n. Male: A-genital lamina and sternogenital shields; B-tectum; C-
gnathosoma and corniculi; D-chelicera, paraxial; E-idem, antiaxial; F-palptrochanter and
palpfemur; G-G’-leg II, femur, genu and tibia antiaxial and ventral; H-coxa II, paraxial.
26 ILINCA JUVARA-BALS & WOJCIECH WITALINSKI
Female: presternal shield with anterior margin serrated; endogynium circular in outline;
its posterior margin protrudes forward to form two prolongations ending beyond the
anterior margin of endogynial sack; endogynial margin and walls with 3-7 denticles
(Fig. 11 D).
Description. Male. Idiosoma markedly elongated, moderately sclerotized.
Dimensions of idiosoma: 615-655 x 390-410 um (L/W factor 1.57-1.64, N=10). Length
of setae: podonotal region 15-20 um, jl=39 um and opisthonotal region 10-13 um.
Ventral side. Genital opening located in a shallow concavity of sternal shield
margin. Genital lamina divided into three lobes: central one bilobate, protruding for-
ward more than lateral ones (Fig. 10 A). Sternogenital region with regular reticulation;
length of sternal setae 39 um. Cuticle around gland pores behind coxae IV not
modified.
Gnathosoma. Tectum trispinate, with all lateral prongs terminated with minute
spines; central prong rounded (Fig. 10 B). Hypostome with conspicuous broad exten-
sion located between slender corniculi. Hypognathal groove with 10 rows of denticles.
Palpcoxal setae pilose, hypostomatic setae simple. Incisions in sclerotized cuticle
behind hypostomatic setae (Fig.10 C).
Chelicera (Fig. 10 D,E). Fixed digit slender; its proximal 3/5 toothless, with
parallel edges; distal part equipped with 4-5 small denticles paraxially, and pilus
dentilis inserted in the vicinity of two more or less pronounced tubercles. Movable digit
curved, bearing 5-7 denticles distally and large tooth medially; dorsal face of digit
weakly sclerotized proximally. An elongated protuberance located paraxially at the
base of spermatotreme. Arthrodial membrane with brush-like process developed
paraxially, synarthrodial membrane rounded.
Pedipalps (Fig. 10 F). Paraxial border of trochanter slightly thickened, visible as
a ridge, ending near seta vl; seta vl thin and barbed along one edge, whereas seta v2
thick and richly pilose. Ventral surface of femur convex.
Legs. Legs I, IN and IV unremarkable. Leg II spurred as follows (Fig. 10 G,G’):
main spur on femur relatively short and rounded, axillary spur rounded and located
close to main spur, both spurs end at the same level. Genu II with conical spur located
distally and extending slightly beyond distal margin. Spur on tibia longer than that on
genu, situated more paraxially and attached to tibia surface over a long distance. Ridge
of 8 denticles and one solitary denticle on coxa II (Fig. 10 H). Measurements: tarsus I =
160-168 um; tarsus IV = 173-184 um.
Female. Idiosoma elongated but highly convex, cuticle moderately sclerotized,
yellow to yellowish-brown. Dimensions of idiosoma: 690-770 x 465-540 um (L/W
factor 1.43-1.52, N=10). Podonotal setae: j1 = 45-46 um, the others 20-39 um; opistho-
notal setae shorter, 10-14 um.
Ventral side. Presternal plates entire, ribbon-like, with denticulate anterior
margin; lateral presternal platelets free. Exopodal shields facing legs I with thickened
anterior edge and connected to sternum via thin cuticle. Reticulation of sternum weakly
pronounced, more visible in anterior region; posterior region with an axially running
granulate strip (Fig. 11 A). Length of sternal setae from 52 um to 59 um.
NEW SPECIES OF HOLOPARASITUS 27
Fıc. 11
Holoparasitus ellipticus sp. n. Female: A-presternal and sternal shields; B-paragynium: C-epi-
gynium; D-endogynium; E-tectum; F-hypognathal groove and corniculus; G-palptrochanter:
H- chelicera, paraxial; I-gland gv2.
Epigynium characterised by sharp lateral spines separated from central apex by
deep concavities; lateral protrusions of subapical epigynial structure extending slightly
beyond epigynium margin (Fig. 11 C). Paragynial shields with oval and slightly
concave metagynial sclerites; thickenings in front of coxae IV absent. Paragynial
28 ILINCA JUVARA-BALS & WOJCIECH WITALINSKI
protrusions “locking” epigynial shield variable, ranging from ones with border
paraxially angular to circular ones (Fig. 11 B). Cup-shaped endogynium circular; its
posterior margin protrudes forward to form two slightly sinuous, tongue-like ribbons
ending beyond anterior margin of endogynium; endogynial margin and walls armed
with several (3-7) denticles, distributed more or less regularly. Endogynial opening
covered with delicate lamina (Fig. 11 D). Opisthogaster with 8 pairs of setae, their
length from 36 um to 48 um. Large, singular gland pores behind coxa IV surrounded by
unmodified cuticle (Fig. 11 I).
Gnathosoma. Tectum trispinate, with long, attenuate median tine (Fig. 11 E).
Hypognathal groove with 10 rows of denticles. Palpcoxal setae pilose, hypostomatic
setae simple, corniculi conical (Fig. 11 F).
Chelicera (Fig. 11 H). Fixed digit with three teeth in front of pilus dentilis;
sometimes central one smaller and located more paraxially. Pilus dentilis protected
partially by truncated cuticular elevation; two teeth and a long, thin cuticular ridge
located behind pilus dentilis. Movable digit with three teeth. Synarthrodial membrane
rounded.
Pedipalps. Paraxial border of trochanter with a slightly thickened ridge, ending
near pectinate seta v2, seta vl pilose (Fig. 11 G). Anterolateral seta of femur spatulate
and pectinate on one edge; genu with anterolateral setae simply spatulate.
Legs. Structure and setation unremarkable. Coxa II provided, on anterolateral
face, with ridge of fine denticles plus one solitary denticle. Measurements: tarsus I =
164-168 um; tarsus IV = 173-183 um.
Discussion. H. ellipticus is very similar in morphology to H. gibber, as well as
to four known species: A. siculus (Berlese, 1906), H. mallorcae Juvara-Bals, 1975,
H. maritimus Hyatt, 1987 and H. lawrencei, Hyatt, 1987. These six species likely
form a monophyletic species-group, which we name mallorcae group with charac-
teristics listed below.
The males can be distinguished mainly due to: sternogenital shield without
excipulum; hypostome (hypostomatic setae on distinct piece of cuticle, hypostome
more or less evidently extended between corniculi); leg II (spur on femur relatively
short).
The females share similar features for: presternal plates (anterior margin serra-
ted, lateral platelets free); endogynium (cup-shaped, circular, with two prolongations
from posterior margin); structure of genital shields ( paragynia devoid of thickening in
front of coxa IV, similar pattern of subapical epigynial structure); structure of chelicera
(3 and 5 denticles on movable and fixed digits, respectively).
Differential diagnosis of the Holoparasitus mallorcae species-group.
Male:
le corniculi indented, tectum with 3 prongs; mobile digit of chelicera with 4-6
denticles; fixed digit denticulated: H. mallorcae Juvara-Bals, 1975
corniculi indented, tectum with | prong; mobile digit of chelicera curved,
enlarged in its upper third with 6 small denticles; fixed digit slender, truncate
apically, with 2-3 denticles on inner margin: H. siculus (Berlese, 1905)
D
NEW SPECIES OF HOLOPARASITUS 29
(59)
corniculi indented, tectum triangular; mobile digit of chelicera with 5 small
denticles distally and 1 larger proximally; fixed digit slightly curved apically
with 7-8 denticles : H. maritimus Hyatt, 1987
4. corniculi conical; mobile digit of chelicera with 6-7 denticles, its dorsal margin
weakly sclerotized proximally, sclerotized microsclerite near basis of spermato-
trema; fixed digit slender with one teeth and 7 denticles around pilus dentilis :
H. ellipticus sp. n.
I corniculi conical; mobile digit of chelicera with | tooth on internal edge and
with one large protuberance on external edge of digit; fixed digit slender with 4
denticles and little tooth around pilus dentilis: H. gibber sp. n.
6. corniculi conical, mobile digit of chelicera with 1-2 teeth on internal edge;fixed
digit edentate, finger-like: H. lawrencei Hyatt, 1987
Female:
Ik apex of epigynium elongated: A. siculus (Berlese, 1905)
2. apex of epigynium normal; endogynium without teeth, endogynial margin
prolongations arcuate: H. maritimus Hyatt, 1987
3: apex of epigynium normal; endogynium without teeth, endogynial margin
prolongations straight: H. mallorcae Juvara-Bals, 1975
4. apex of epigynium normal; endogynium with two teeth, prolongations sinuous:
H. gibber sp. n.
> apex of epigynium normal; endogynium with several teeth, prolongations
sinuous: A. ellipticus sp. n.
6. apex of epigynium enlarged with little tip medially; endogynium with many
internal teeth, prolongations curved, usually with denticles or corrugations at the
end: A. lawrencei Hyatt, 1987
ACKNOWLEDGEMENTS
We express our sincere appreciation to the following persons for providing us
with the material used in the descriptions: Dr. C. Athias-Henriot, Miremont-Haut
Mauzens, France; Drs M. Castagnoli, F. Pegazzano and R. Nannelli, Experimental
Institute of Agricultural Zoology, Florence, Italy; Prof. Dr. R. Dallai, University of
Siena, Siena, Italy; Prof. Dr. V. Mahnert, Dr. B. Hauser, Museum of Natural History,
Geneva, Switzerland, Dr. L. Tiefenbacher, Munich, Germany. We are also indebted to
Drs F. Pegazzano and M. Castagnoli for some observations on type material in Berlese
Acaroteca. For reviewing the manuscript and for their helpful comments, we kindly
thank Dr. E. Lindquist, Centre for Land and Biological Resources Research, Ottawa,
Canada and Dr. I. Löbl, Museum of Natural History, Geneva.
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30 ILINCA JUVARA-BALS & WOJCIECH WITALINSKI
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SCHMÖLZER, K. 1991. Landmilben aus Kärnten I. (Acarina, Parasitiformes). Carinthia II, 181:
343-358.
SCHMOLZER, K. 1995a. Einige neue Landmilben aus Südkärnten (Acari,Parasitiformis) Atti del
Museo Civico di Storia Naturale di Trieste 46: 99-112.
SCHMÖLZER, K. 1995b. Landmilben aus den östlichen Karawanken (Acarina, Parasitiformis)
Carinthia II, 185: 499-512.
Turk, F. A. 1953. A synonymic catalogue of British Acari. Annals and Magazine of Natural
History 6: 1-26, 81-99.
VAN DER HAAMMEN, L. 1980. Glossary of Acarological Terms. Vol.I. General Terminology,
W. Junk Publishers, The Hague, 244 pp.
VINNIK, O. M. 1994. A new species of the genus Holoparasitus (Mesostigmata, Parasitiformes)
from the Crimea. Vestnik Zoologii Kiev 2: 82-84.
VITZTHUM, H. v. 1923. Acarologische Beobachtungen. 7. Reihe. Archiv für Naturgeschichte
89A: 97-181.
WITALINSKI, W. 1994a. Holoparasitus (Holoparasitus) dallaii sp. n., a new gamasid mite from
Sardinia, Italy (Acarı: Pergamasidae). International Journal of Acarology 19: 349-354.
WITALINSKI, W. 1994b. Two new Holoparasitus species from Italy and Poland (Acari: Gamasida:
Pergamasidae). Genus 5: 215-222.
REVUE SUISSE DE ZOOLOGIE 107 (1): 31-48; mars 2000
Three new species of Geodia Lamarck, 1815
(Porifera, Demospongiae)
from the bathyal depths off Brazilian coast, Southwestern Atlantic
Carla Maria Menegola da SILVA! & Beatriz MOTHES?
| Universidade de Säo Paulo, Museu de Ciéncias Naturais, Fundaçäo Zoobotänica do
Rio Grande do Sul, Av. Salvador Franga, 1427, 90690.000, Porto Alegre, RS, Brasil.
E-mail: silva@ portoweb.com.br
2 Museu de Ciéncias Naturais, Fundacäo Zoobotanica do Rio Grande do Sul,
Caixa Postal 1188, 90001-970, Porto Alegre, RS, Brasil.
E-mail: bmothes @ portoweb.com.br
Three new species of Geodia Lamarck, 1815 (Porifera, Demospongiae)
from the bathyal depths off Brazilian coast, Southwestern Atlantic. -
This work comprises a taxonomic study of tetractinellid sponges from a
poorly known region in the southwestern Atlantic, off the Rio Grande do
Sul State coast, Brazil (31°05’—32°00’S/49°31’-50°00’W). Samples were
collected by R/V “Atlantico Sul” of Fundagäo Universidade do Rio Grande
in a continental slope survey (“Projeto Talude”) and R/V “Prof. W. Bes-
nard”, of Instituto Oceanografico da Universidade de Sao Paulo/Group for
the Development of the Fisheries Industry, in Rio Grande do Sul I Cruise.
Three new species are described: Geodia australis, G. splendida and
G. riograndensis.
Key-words: Porifera - Geodia - Rio Grande do Sul - Brazil - taxonomy -
continental slope.
INTRODUCTION
Six species of Geodia are registered for the Brazilian coast: Geodia gibberosa
Lamarck, 1815 (Laubenfels, 1956), Geodia neptuni (Sollas, 1886, 1888 as Synops
neptuni; Mothes, 1996), Geodia tylastra Boury-Esnault, 1973 (Boury-Esnault, 1973),
Geodia papyracea Hechtel, 1976 (Hechtel, 1976), Geodia corticostylifera Hajdu et al.,
1992 (Hajdu ef al., 1992) and Geodia glariosa (Sollas, 1886, 1888 and Volkmer-Ribeiro
& Mothes-de-Moraes, 1975 as Cydonium glariosus).
The present paper describes three new species dredged off Rio Grande do Sul
State coast (31°05’-32°00°S/49°31’-50°00’W) (fig. 1), in the slope region (207 to
520 m depth), during oceanographic expeditions carried out by R/V “Atlantico Sul”, of
Manuscript accepted 12.11.1999
32 CARLA MARIA MENEGOLA DA SILVA & BEATRIZ MOTHES
Fundacäo Universidade do Rio Grande, Projeto Talude; and by R/V “Prof. W. Bes-
nard”, of Instituto Oceanogräfico, Universidade de Säo Paulo, in agreement with
Group for the Development of the Fisheries Industry, in Rio Grande do Sul I Cruise.
The influence of the Subtropical Convergence, with marked seasonal latitu-
dinal displacement, characterizes the southern/southeastern Brazilian continental shelf
and slope regions (23°S-34°S) as a biogeographic transition zone (Mothes, 1996;
Sharp, 1988) between the large neritic areas of Patagonia and tropical Brazil. The
composition and abundance of species, the pelagic structure, the spatial distribution of
communities and their trophic interactions, as well as biological production are
largely controlled by the seasonal dominance of distinct water masses over shelf and
slope. The studied material was collected in the summer period (october to april),
when the influence of Tropical Waters is greatest, though waters of subantarctic
origin may also rise during the summer along the southernmost shelf break regions.
The paratypes of Geodia australis sp. n. were collected between Sarita and Rio
Grande localities, 101 Km off Rio Grande do Sul coast, with temperature of 14,50°C
and salinity 35,76%c. The holotype of this species was collected between Mostardas
and Solidao localities, 58 Km off Rio Grande do Sul coast. Temperature and salinity
data for this sample are not known, as well as for the type-specimens of Geodia
splendida sp.n. and G. riograndensis sp. n.
MATERIAL AND METHODS
The samples are deposited in the Porifera Collection of Museu de Ciéncias
Naturais, Fundaçäo Zoobotanica do Rio Grande do Sul (MCN/ FZB).
Abbreviations used are:
BMNH Natural History Museum, London
MCN/FZB Museu de Ciéncias Naturais, Fundaçäo Zoobotanica do Rio Grande do
Sul, Porto Alegre, Brazil
MHNG Muséum d histoire naturelle Genéve, Switzerland
UFRJ Universidade Federal do Rio de Janeiro, Rio de Janeiro, Brazil
ZMA Zoölogisch Museum Amsterdam, Netherlands
The methodology used to prepare thick sections and dissociated spicules slides
follow Mothes-de-Moraes (1978). Electron micrographs were taken at MCN/FZB
with a Jeol 5200 equipment, with an accelerating voltage of 25 kV and magnifications
varying from 1,500 to 10,000 times. Spicule measurements refer to minimum, mean,
and maximum sizes in micrometers (um) and were obtained by taking 50 measures of
each type of spicule/specimen (unless stated otherwise).
DESCRIPTIONS
Order Astrophorida
Geodiidae Gray, 1867
Geodia Lamarck, 1815
THREE NEW SPECIES OF GEODIA 33
40°W
South America
Fic. 1
Map of South American Coast with the Rio Grande do Sul State Coast in detail, showing the
geographic distribution of Geodia australis sp. n. (MW), G. splendida sp. n. (A) and G. riogran-
densis sp. n. (D)).
Type-species: Geodia gibberosa Lamarck, 1815
Definition: Geodiidae with afferent and efferent aquiferous system indepen-
dently, with well developed and large subectosomal spaces. Megascleres triaenes.
Microscleres sterrasters and euasters of different types. Sterrasters varying from
flattened or globose young forms, smooth with many spherical spaces or provided of
conical and short rays, with blunt or strongiliform ends sometimes presenting small
distal holes, to globose forms provided of star or rosette like microspinature at the distal
end (adapted from Desqueyroux-Fatindez & Van Soest, 1997).
Geodia australis sp. n. Figs 1, 2, 5, 7, 11-26
Material: Holotype MCN 330, R/V “Prof. W. Besnard”, off Rio Grande do Sul State
coast, St. 458 (33°29’S/50°44’ W), 9/XII/1968, 207 m, rocky substrate; Paratype MHNG-INVE
34 CARLA MARIA MENEGOLA DA SILVA & BEATRIZ MOTHES
26564 (schizoparatype-slides deposited in the MCN 331), same data of the holotype; Paratype
ZMA POR13418 (schizoparatype-slides deposited in the MCN 332), R/V “Prof. W. Besnard”,
off Rio Grande do Sul State coast, St. 444 (31°31’00”S/49°47’00”W), 6/XII/1968, 284 m, rocky
substrate.
Description: Spherical sponge (fig. 2) (diameter 3.8 cm x 3.1 cm, height 2.8
cm). Hispid surface, with slight brushes of oxeas (fig. 7); small openings are observed
in some points of the surface, which could not be differentiated in ostia or oscula.
Colour in spirit grayish-white; compressible consistency. The sponges are associated
with polychaets and corals.
Skeleton: Ectosomal (fig. 5): cortex (0.5 mm thick) with spherasters of variable
sizes, over several layers of sterrasters ended at the cladome-layer of the dichotriaenes,
plagiotriaenes and protriaenes; scattered small oxeas, strewn at random, are also
observed.
Choanosomal (fig. 5): formed by dichotriaenes, plagiotriaenes, protriaenes and
large oxeas, the last ones preferentially arranged in radial bundles, forming a right angle
with the cortex. Besides the triaenes and oxeas, sterrasters, spherasters and oxyasters
are abundantly spread.
Spicules: Oxeas I (fig. 12): fusiform, straight or slightly curved, with pointed or
blunt ends; some with mucronate ends; length 1403-2285. 7-3818/24-31.0-40 um.
Oxeas II (fig. 13): fusiform, with gradually pointed ends; length 181-275. /-
418/3.2-4.8-6.9 um.
Dichotriaenes (fig. 11): conical rhabdom, with gradually pointed or strongyloid
end. Cladi are first curved upwards and then slightly downwards; rhabdome 1012-
2246.0-3565/33-61.0-86 um: cladome 333-553.8-703 um, protocladi 161-/92./-
238/29-38.7-48 um; deuterocladi 76-/07.2-143 um.
Protriaenes (fig. 17): conical, thin rhabdome, with gradually pointed or blunt
end; cladi with thin ends, sometimes provided with a constriction; rhabdome 1334-
3143.9-5865/4.6-10.4-23 um; cladome 86-/54.2-276 um, cladi 67-156.4-276/4.6-8.2-
14 um.
Plagiotriaenes (fig. 16): rare (N=5), rhabdome conical, straight and thin, with
gradually pointed end; cladı with thin ends; rhabdome 828-1909/19-28.5 um; cladome
105-219 um; cladi 51-131/11.5-18.4 um.
Anatriaenes (fig. 14): rhabdome conical and thin, with pointed or strongyloid
end; cladi with pointed ends, sometimes marked by a constriction; rhabdome 1150-
3450.0-6140/4.8-14.2-24 um; cladome 67-/30.9-190 um, cladi 48-//0.6-181 um.
Some rares (N=5), slender and smaller anatriaenes (fig. 15) can be observed: rhabdome
© 323-423/3.4-4.6 um; cladome 6.9-9.2 um; cladi 2.3-4.6/1.1-1.8 um.
Sterrasters (figs. 18-20): oval or more rarely spherical, the young scleres with
microspined surface in the shape of pointed cones and in the shape of a star in adult
scleres; diameter 266-3 /4.5-352/190-253.6-295 um.
Spherasters (fig. 21): spherical with well delimited center, variable size and
short conical spines with blunt ends; in the choanosome, the largest spherasters can be
taken for young sterrasters, differing by the shape of the microspines; diameter 20-26.3-
32 um.
THREE NEW SPECIES OF GEODIA 35
FIGs 2-4
Habit: 2, Geodia australis sp. n.; 3, G. splendida sp. n.; 3, G. riograndensis sp. n.-inner view in
transversal section. Scale = 2 cm.
36 CARLA MARIA MENEGOLA DA SILVA & BEATRIZ MOTHES
Oxyasters I (fig. 22): small and smooth center; 4 to 7 long, conical and micro-
spined rays; total diameter 35-43.2-52 um, diameter of the centrum 2.3-4.3-6.9 um,
rays 9-16.9-25/1.5-2.3-3.5 um.
Oxyasters II (fig. 23): small and smooth center: 8 to 11 long, conical and micro-
spined rays; total diameter 16-23.7-32 um, diameter of the centrum 2.3-4.2-5.8 um,
rays 6.9-/0.0-13.8/2.3 um.
Spheroxyasters (fig. 26): discrete center; 15 to 22 short , conical and scarcely
microspined rays; total diameter 14-/6./-20 um, diameter of the centrum 4.6-5.7-6.9
Ria clear And smooth; 7 to 14 short, blunt rays, with conical
A total diameter 4.6-7.0-9.2 um, diameter of the centrum 2.3-2.8-3.4 um,
rays 1.8-2.3-2.8/<1.0 um.
Etymology: The specific name refers to the type-locality, off Brazil’s southern
region [Latin Bai australis = southern: south].
Remarks: The samples utilized in this description were identified by Mothes-de-
Moraes (1978) as Geodia eosaster (Sollas, 1886).
Hajdu et al. (1992) advanced the idea that G. eosaster sensu Mothes-de-Moraes
(1978) could be a new species due to its widely disjunct distribution when compared to
the original record of G. eosaster from Australia (Sollas, 1886; 1888). Our reexa-
mination of Mothes-de-Moraes (1978) specimens, when compared with the syntype of
G. eosaster [BMNH 1889.1.1.87], revealed several spicular micrometric distinctions
and the SEM analysis confirmed the presence of adittional categories of megascleres
and microscleres. We thus described them as Geodia australis sp. n.
Both species share the presence of dichotriaenes, protriaenes, spherasters and
strongylasters, but they can be distinguished by the occurrence of rare plagiotriaenes,
oval shaped sterrasters, and of a second type of oxyaster in the new species.
From the Tropical western Atlantic records of Geodia, the new species appears
closest to G. spherastrea Lévi, 1964, from deep-waters off Puerto Rico, at 2840 m
depth. Both species share the presence of dichotriaenes, protriaenes, anatriaenes, spher-
asters and strongylasters, but can be distinguished by the occurrence of rare plagio-
triaenes, oxyasters and oval shaped sterrasters in the new species.
The “somal” spherasters of Geodia eosaster (Sollas, 1886) and the chiasters
[=strongylasters sensu Boury-Esnault & Riitzler, 1997] of G. spherastrea (Lévi, 1964),
correspond to the spherostrongylasters here described for G. australis sp. n. The term
was coined for cases where the width of the aster’s centrum exceeds 1/3 of the
microscleres total diameter.
Geodia splendida sp. n. Figs 1, 3, 6, 8, 27-39
Material: Holotype MCN 2355 (schizoholotype MHNG-INVE 26565), R/V “Atlantico
Sul”, off Rio Grande do Sul State coast, St. 10 (32°00°S/50°00’W), 10-X-1991, 520 m, rocky
bottom.
Description: Globose sponge (fig. 3), diameter 27 cm, height 24 cm. Surface
smooth to the touch; single oscule, apical, central, elypsoidal, diameter 3.5/3 cm, at the
THREE NEW SPECIES OF GEODIA 37
Serum 2353581
Fics 5-8
Skeleton: 5, 6.Skeletal arrangement. 5, Geodia australis sp. n.; 6. G. splendida sp. n.; 7, 8.
Sponge surface : 7, G. australis sp. n.; 8, G. splendida sp. n.
38 CARLA MARIA MENEGOLA DA SILVA & BEATRIZ MOTHES
159182
Fics 9-10
Geodia riograndensis sp. n.: 9, skeletal architecture: 10, oxeas protracting at the sponge surface.
THREE NEW SPECIES OF GEODIA 39
Il
12 8
Mi
Fics 11-17
Megascleres of Geodia australis sp. n.: 11, dichotriaene; 12, oxeas I; 13, oxea II; 14, anatriaene:
15. smaller anatriaene; 16, plagiotriaene; 17, protriaene. Scales = 100 um.
——
17
40 CARLA MARIA MENEGOLA DA SILVA & BEATRIZ MOTHES
terminal part of a cylindrical channel (length 12 cm), in the inner lateral walls of which
the exhalant channel openings can be observed; pores not visible. Preserved material of
violet colour and hard consistency.
Skeleton: Ectosomal (fig. 6): cortex made up of several overlaping layers of
sterrasters (0.6-0.7 cm) and the discrete protraction of robust oxeas and some rare
styloid forms (fig. 8); the cladomes of the orthotriaenes are placed parallel to the
sponge surface just below the cortex.
Choanosomal (fig. 6): formed by the rhabdoms of the orthotriaenes, perpendi-
cular to the surface and, among them, long oxeas, randomly distributed and rare
sterrasters.
Spicules: Oxeas I (fig. 27): straight or slightly bent, robust, with ends blunt or
gradually pointed, length 2254-2681.0-3151/ width 28-44.2-62 um.
Oxeas II (fig. 30): straight or slightly curved, with gradually pointed or mucro-
nate ends; some thinner scleres have blunt ends. Length 228-432.7-684 um, width 5.7-
10.7-19 um.
Orthotriaenes (figs 28, 29): straigth; cladı straight or bent downward at their
distal portion, with blunt or gradually sharpening ends; length 3266-3689.0-4094 um,
width 104-///.6-120 um, cladome diameter 920-//65.7-1495 um.; cladi length 437-
589.8-759 um, cladi width at the base 85-/00.8-113 um.
Sterrasters (figs 31-36): spherical or ellipsoidal, with conspicuous hilum, micro-
spined at the outer portion (figs 34, 35); surface with rounded holes (figs. 31, 32) or
conical rays (fig. 33) in young forms; or provided with regular microspinature in star
shape in adult scleres (figs 34-36); diameter 95-148.2-171/86-//9.7-152 um.
Oxyasters I (fig. 37): 4 to 8 microspined rays all along their length; total
diameter 78-/00./-131 um, center 6.9-9.2-13.8 um, rays 35-46.9-62/2.8-4.7-6.9 um.
Oxyasters II (fig. 38): 4 to 9 rays provided with conical microspines all along
their length, total diameter 12-/7.9-23 um, center 1.6-2.3-3.4 um, rays length 4.6-7.9-
11.5 um, rays width < | um.
Spherostrongylasters (fig. 39): 5 to 11 microspined rays all along their length,
with blunt ends, diameter 5.5-7.4-9.9 um, rays 1.1-/.5-2.2/<1.0-1.5 um.
Etymology: The specific name refers to the beauty and large size of the
specimen. [Latin word splendidus = magnific]
Remarks: Geodia splendida sp. n. is close to Geodia corticostylifera Hajdu et
al., 1992 [Holotype UFRJ POR 3098 and Paratype UFRJ POR 3714, examined] by the
shared presence of oxeas, orthotriaenes and oxyasters. They both differ nevertheless by
the presence of an additional category of styles instead of oxeas in G. corticostylifera
and of microscleres of the spheroxyaster and strongylospheraster types in G. splendida
sp. n.
Geodia riograndensis sp. n. Figs 1, 4, 9, 10, 40-58
Material: Holotype MCN 1591 (schizoholotype MHNG-INVE 26566),“R/V Atlantico
Sul”, off Rio Grande do Sul State coast, St. 2-26 (31°05’S/49°31’W), 15-II-1987, 300 m, rocky
substrate; Paratype MCN 3452, R/V . “Atlantico Sul”, off Rio Grande do Sul State coast, St. 1-5
(32°24°55”S/50°14°85”W), 30/1V/1986, 200 m, rocky substrate.
THREE NEW SPECIES OF GEODIA 4]
25V x1.,000
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25kU *1-560 18mm 633221 pus Sue 033228
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Fics 18-26
Microscleres of Geodia australis Sp. n.: 18, young sterraster; 19, adult sterraster surface; 20, adult
sterraster with hilum; 21, spheraster and spherostrongylaster (arrow); 22, oxyaster I; 23, oxyaster
II; 24, spherostrongylaster with conical tips; 25, spherostrongylaster; 26, spheroxyaster.
42 CARLA MARIA MENEGOLA DA SILVA & BEATRIZ MOTHES
28
29
30
Fics 27-30
Megascleres of Geodia splendida sp. n.: 27, oxea I; 28, orthotriaene; 29, orthotriaene cladome in
apical view; 30, oxeas II. Scale = 500 um.
THREE NEW SPECIES OF GEODIA 43
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2355514
Fics 31-39
Microscleres of Geodia splendida sp. n.: 31, 33 sterraster developmental stage; 32, sterraster
developmental stage surface; 34, adult sterraster; 35, sterraster surface with hilum; 36, adult
sterraster showing microspinature details; 37, oxyaster I; 38, oxyaster II: 39, spherostrongylaster
developmental stage (arrow) and grown spherostrongylaster.
44 CARLA MARIA MENEGOLA DA SILVA & BEATRIZ MOTHES
Description: Subglobose fragment (fig. 4), diameter 10.6 cm, height 7.5 cm.
Hispid surface, with slight brushes of oxeas (0.2-0.5 mm) (fig. 10); central oscule
(diameter 11 mm), raising above the surface (3 mm), at the terminal part of a cylindric
channel (length 44 mm), in the inner lateral wall of which the opening of the exhallant
channels can be observed; pores were not observed. Preserved material with beige
colour and firm consistency.
Skeleton: Ectosomal (fig. 9): Cortex made up by large quantities of sterrasters in
regular overlapping layers, throughout which, small and thin oxeas are found and, more
rarely, very long oxeas with filiform ends, both projecting slightly above the sponge
surface. In the subcortical area orthotriaenes and additional categories of triaenes are
distributed slant or perpendicular to the surface, the latter being hardly observed.
Choanosomal (fig. 9): formed by tracts of oxeas perpendicular to the surface
and rare sterrasters of random distribution.
Spicules: Oxeas I (fig. 42): thin, sinuous, with gradually pointed ends, length
2415-5720.5-8464 um, width: 12-/7.5-23 um.
Oxeas II (fig. 43): robust, straight or slightly curved, with blunt or mucronate
ends. Some scleres thinner, with one of the ends presenting lateral conical expansions,
from which one of the sclere sides becomes gradually thinner, length 1610-2082.0-2726
um, width 21-35.2-46 um.
Oxeas III (fig. 44): straight or slightly curved, with gradually pointed ends,
length 247-486.6-741 um, width 5.7-8.9-11.4 um.
Orthotriaenes (figs 40, 41): straight rhabd with end conical or sharpening gra-
dually; straight or downwards cladi with conical or strongyliform distal ends, some-
times bi- or trifurcate, length 1725-28/9.8-3675 um, width 44-66.5-92 um, cladome
diameter 575-775.3-989 um, cladi length 253-365.4-437 um, cladi thickness at the base
32-55.0-69 um.
Anatriaenes (fig. 50): rare (N=6); straight rhabd with gradually pointed ends,
cladi with gradually pointed or strongyliform ends, length 4501 um, width 9.5-12.3 (m,
cladome diameter 33-67 um, cladi length 19-38 um, cladi thickness at the base 6.6-9.5
um.
Plagiotriaenes (fig. 49): rare (N=4); straight rhabd with gradually pointed end:
cladi gradually pointed, length 1495-1886 um, width 28-39 um, cladome diameter 460-
506 um, cladi length: 230-253 um, cladi thickness at the base 23-37 um.
Protriaenes (figs 47, 48): rare (N=6), straight or slightly curved rhabd, with blunt
or abruptly pointed end, length 3030-5282 um, width: 9.5-19 um; cladome diameter
95-204.2 um, cladi length 62-124 um, cladi thickness at the base 8.5-14.2 um.
Anamesotriaenes (fig. 51). rare (N=5); straight rhabdome with abruptly pointed
or strongyliform end; cladi with conical or strongyliform ends, length 5938-7581 um,
width 9.5- 14.2 um, cladome diemeter 49-67 um, cladi length 19-38 um, cladi
thickness at the base 6.7- 14.2 um.
Promesotriaenes (fig. 45): rare (N=3); straight rhabdome with gradually pointed
end; cladi with gradually pointed or strongyliform end, length 2484-3404 um, width
9.5-19 um, cladome diameter 105-190 um, cladi length 52-105 um, cladi thickness at
the base 9.5-16.1 um.
THREE NEW SPECIES OF GEODIA 45
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47 |
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48
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Fics 40-51
Megascleres of Geodia riograndensis sp. n.: 40, orthotriaene; 41, orthotriaene cladome in apical
view; 42, oxea I; 43, oxeas II; 44, oxeas III; 45, promesotriaene; 46, diaene; 47, protriaene; 48,
protriaene basal extremity; 49, plagiotriaene; 50, anatriaene; 51, anamesotriaene. Scales =
200 um.
46 CARLA MARIA MENEGOLA DA SILVA & BEATRIZ MOTHES
25kU x3,See Sem 159107
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FIGS 52-58
Microscleres of Geodia riograndensis sp. n.: 52, sterraster, oxyaster I and oxyaster II (arrow); 53,
sterraster surface with hilum; 54, oxyaster I and II (arrow); 55, oxyaster II e spherostrongylaster
(arrow); 56, strongylaster and spherostrongylaster (arrow); 57, strongylaster; 58, strongylaster
(arrow) and spherostrongylaster.
THREE NEW SPECIES OF GEODIA 47
Diaenes (fig. 46) : rare (N=2); straight or slightly sinuous rhabdome, with
gradually pointed end, length 184 um, width 17 um, cladome diameter 200 um, cladi
length 157 um, cladi thickness at the base 13.3 um.
Sterrasters (figs 52, 53): spherical or oval; hilum spherical and conspicuous;
surface provided with irregular microspines in form of a rosette, sometimes absent in
the region around the hilum: diameter 57-98. /-124 um/48-89.9-114 um.
Oxyasters I (figs 52, 54): 3 to 8 microspined rays distributed along its whole
length, diameter 64-86,4-117 um, center: 5.7-8.3-11.5 um, rays length 30-4/.9-58 um,
rays width 2.3-3.8-4.6 um.
Oxyasters II (figs 52, 54, 55): 4 to 9 microspined rays at the distal portion or,
more rarely, all along their length, diameter 14-20.2-30 um, center 1.8-3.0-4.6 um, rays
length 4.6-9.0-13.8 um, rays width 1.1-/.6-2.3 um.
Strongylasters (figs 56-58) - varying to spherostrongylasters (figs 55, 56, 58). 5
to 13 rays with strongyliform or truncate ends, microspined all along their length or,
more rarely, at the distal half, diameter 4.6-8.4-13.2 um, center 1.6-3.2-4.6 um, rays
length: <1.0-3.0-4.6/< 1.0 um.
Etymology: The specific name refers to the Rio Grande do Sul State coast, off
which the sponges were collected.
Remarks: When compared with other species of Geodia from the Brazilian
coast, Geodia riograndensis sp. n. is close to G. glariosa Sollas, 1886 [Syntype BMNH
1889.1.1.86] by the shared occurrence of oxeas, ortho-, pro- and anatrienes as mega-
scleres, differing, however, for presenting additional categories of megascleres, as three
categories of oxeas, plagiotriaenes, anamesotriaenes, promesodiaenes, promesotriaenes
and diaenes; and of microscleres, as oxyasters, strongylasters and strongylospherasters.
ACKNOWLEDGEMENTS
We are thankful to Fundaçäo de Amparo a Pesquisa do Estado do Rio Grande do
Sul (FAPERGS) for granting the Master’s Newly-Graduated Scholarship to the senior
author (Proc. n° 96/60364-1) in the first part of the work (study of Geodia riogran-
densis sp. n. and G. splendida sp. n.) and Aid to the junior author (Proc. n° 95/ 0728.3);
to Fundaçäo de Amparo a Pesquisa do Estado de Sao Paulo (FAPESP) for granting the
Doctoral’s Scolarship (study of Geodia australis sp. n. and conclusion of the paper) to
the senior author (Proc. n° 98/00797-4); to Dr. Ricardo R. Capitoli (FURG), for the
donation of the specimens studied; to Dr. Clare Valentine (BMNH) and Dr. Guilherme
Muricy (UFRJ), for lending the type-materials intended for comparative studies; to the
Museum of natural history of Geneva (MHNG) for cooperation; to Dr. Eduardo Hajdu
(MNRJ) and M. Sc. Cléa B. Lerner (USP), for their critical reading and suggestions for
the paper; to the technicians Cleodir J. Mansan and Marcia Spadoni (MCN), for the
scanning electron microscope photos and to Mrs. Rejane Rosa (MCN) for the final
scleres drawings and map.
48 CARLA MARIA MENEGOLA DA SILVA & BEATRIZ MOTHES
REFERENCES
BOURY-ESNAULT, N. 1973. Campagne de la Calypso au large des côtes atlantiques de 1° Amérique
du Sud (1961-1962). I, 29. Spongiaires. Results of scientific Campaign of Calypso 10:
263-295.
Boury-ESNAULT, N. & RÜTZLER, K. 1997. Thesaurus of Sponge Morphology. Smithsonian
Contributions to Zoology 596: 1-55.
DESQUEYROUX-FAUNDEZ, R. & VAN SOEST, R. W. M. 1997. Shallow waters Demosponges of the
Galäpagos Islands. Revue suisse de Zoologie 104 (2): 379-467.
HaJpu, E. C. M, Muricy, G., CusToDIo, M., Russo, C. & PEIXINHO, S. 1992. Geodia
corticostylifera (Demospongiae, Porifera) New Astrophorid from the brazilian coast
(Southwestern Atlantic). Bulletin of Marine Science 51 (2): 204-217.
HECHTEL, G. J. 1976. Zoogeography of Brazilian Marine Demospongiae (pp. 237-259). In:
Harrison, F. W. & Cowden, R. R. (eds). Aspects of Sponge Biology.
LAUBENFELS, M. W. DE. 1956. Preliminary discussion of the sponges of Brazil. Boletim do
Instituto Oceanografico 1: 1-4.
Levi, C. 1964. Spongiaires des zones bathyale, abyssale et hadale (pp. 63-112). In: Torben Wolff
(ed.). Galathea Report, Scientific results of the Danish Deep-Sea Expedition Round the
World, 1950-1952.
MOTHES, B. 1996. Esponjas da Plataforma Continental Norte e Nordeste do Brasil. Unpublished
thesis, Universidade de Sao Paulo, Instituto de Biociéncias, Sao Paulo, SP, 230 pp.
MOTHES-DE-MORAES, B. 1978. Esponjas tetraxonidas do litoral sul-brasileiro: II. Material cole-
tado pelo N/Oc. “Prof. W. Besnard” durante o Programa RS. Boletim do Instituto
Oceanografico 27 (2): 57-78.
SHARP, G. D. 1988. Fish populations and fisheries (pp. 155-202). In: H. Postma & J.J. Zijlstra
(eds). Continental Shelves. Ecosystems of the World, vol. 27. Elsevier, Amsterdam.
SOLLAS, W. J. 1886. Preliminary account of the tetractinellid sponges dredged by H.M.S.
“Challenger”, 1872-76. Part I. The Choristida. Scientific Proceedings of the Royal Dublin
Society 5 (4): 177-199.
SOLLAS, W. J. 1888. Report on the Tetractinellida collected by the H. M. S. “Challenger”
during the years 1873-1876. Report on the scientific results of the voyage of H. M. S.
“Challenger” Zool. 25: xi-clxvi + 1-457 pp.
VOLKMER-RIBEIRO, C. & Mothes-de-Moraes, B. 1975. Esponjas tetraxonidas do litoral sul-brasi-
leiro. I - Redescricào de Cydonium glariosus Sollas, 1886 e Erylus formosus Sollas,
1886. Iheringia, ser. Zool. (47): 3-22.
REVUE SUISSE DE ZOOLOGIE 107 (1): 49-79; mars 2000
Oribatids from Switzerland III (Acari: Oribatida: Oppiidae 1
and Quadroppiidae).
(Acarologica Genavensia XCIII)
Sandor MAHUNKA & Luise MAHUNKA-Papp
Zoological Department, Hungarian Natural History Museum, Baross utca 13,
H-1088 Budapest, Hungary.
Oribatids from Switzerland III (Acari: Oribatida: Oppiidae 1 and
Quadroppiidae). (Acarologica Genavensia XCHI). — Oppioid oribatids
taken from soil samples in Switzerland are recorded. There are 15 species,
of which 6 are new for science and one of them also represents a new
genus: Paramedioppia gen. n. (Oppiidae). The following 5 new combina-
tions are proposed: Berniniella conjuncta (Strentzke) comb. n. et stat. n. =
Oppia sigma conjuncta Strentzke, 1951; Berniniella hauseri (Mahunka)
comb. n. = Oppia hauseri Mahunka, 1974; Berniniella serratirostris
(Golosova) comb. n. = Oppia serratirostris Golosova, 1970; Dissorhina
signata (Schwalbe) comb. n. = Oppia signata Schwalbe, 1989; Lauroppia
maritima (Willmann) comb. n. = Oppia maritima Willmann, 1929; 1 spe-
cies is placed in synonymy: Oppiella rafalskii Optotna & Rajski, 1983 =
Berniniella hauseri (Mahunka, 1974): syn. n. In addition 2 species (still not
recorded for Switzerland) are discussed, in this way a total of 17 oppioid
species are considered. Morphological and distributional data of 11 species
are provided and the nature of relationships and some additional morpholo-
gical characters are discussed.
Key-words: Acari - Oribatida - Oppiidae - Quadroppiidae - taxonomy -
new species - new genus - new combinations - Switzerland.
INTRODUCTION
Our revision work regarding the oribatids of Switzerland, which will eventually
be incorporated into a book, has several times been mentioned (e.g. Mahunka, 1993,
1996a). For the very simple reason that at least another 4-5 years will have to elapse
until the appearence of this book, we believe that all the taxonomic novelties and
specific faunistic data should be published in order to complement the ever increasing
number of taxonomic and zoogeographic researches. This time we propose to discuss
a part of the available information gained in connection with species belonging to the
superfamily of Oppioidea.
Manuscript accepted 13.01.1999
50 SANDOR MAHUNKA & LUISE MAHUNKA-PAPP
We should stress that, in referring to the taxon superfamily, we adopt the
system incorporating e.g. the following families: Oppiidae, Quadroppiidae and
Suctobelbidae. In this paper we discuss especies belonging to the families Oppiidae
Grandjean, 1951 and Quadroppiidae Balogh, 1983. The subfamilial division within
the family Oppiidae seems quite unsuitable for further subdivisions, especially since
there are many genera (see Mahunka, 1999).
List of localities
GE-4 = Switzerland: Genève: Frontenex, pieds souches chênes; 14.VII.1980; leg. C. Besuchet
— (33).
GR-6 = Switzerland: Graubünden: Landquart, pieds aulne (Alnus); 26.1X.1983; leg. C. Besu-
chet — (118).
GR-8 = Switzerland: Graubiinden: Samnaun, alpine Wiesen mit Rhododendron, Gesiebe,
2050m; 26.VIIL. 1968; leg. C. Besuchet — (26).
GR-9 = Switzerland: Graubünden: Santa Maria — Paß Umbrail, Gesiebe, 2000m; 5.VIII.1974;
leg. C. Besuchet — (37).
GR-10= Switzerland: Graubünden: Untervaz b. Chur, mousses: 29.IX.1983; leg C. Besuchet —
(126).
LU-I = Switzerland: Luzern: Eigenthal, pres Eigenthal (village) Forenmoos, Sphagnum,
970m.; 2.VIIL. 1996; leg. C. Besuchet — (108).
NW-1 = Switzerland: Nidwald: Musenalp, oberhalb Niederrickenbach, mousses dans lappiaz,
1800m; leg. C. Besuchet — (113).
NW-2 = Switzerland: Nidwald: Pilatus, tamisage rhododendron, 1800m; 14.VI.1984; leg.
I. Löbl — (123).
Switzerland: Nidwald: Trübsee, mousses près source, 1800m; 8.1X.1997; leg.
C. Besuchet — (127).
SO-5 = Switzerland: Solothurn: Schottwill, Bucheggberg, Rindenmoos von lebendem Baum;
27.1X.1987; leg. S. Mahunka & L. Mahunka-Papp — (47).
SZ-3 = Switzerland: Schwyz: Pragelpaß, mousses sapins, 1650m; 25.VIII.1983; leg. I. Lobl —
(116).
TI-5 = Switzerland: Tessin: Monadello - Moneto, im faulenden Laub, 850m; 23.IV.1992;
leg. C. Besuchet — (91).
TI-9 = Switzerland: Tessin: Nufenen-Paß, Boden, dürres Laub und Baummulm aus einem
Lärchenwald; 15.VI.1979: leg. S. Mahunka & L. Mahunka-Papp - (18).
TI-11 = Switzerland: Tessin: Rancate, forêt de chätaigniers, tamisages; 7.1X.1965; leg.
C. Besuchet — (25).
VS-4 = Switzerland: Valais: Daubensee, mousses et herbes, 2200m; 11.VIII.1980; leg.
C. Besuchet — (32).
VS-8 = Switzerland: Valais: Forêt de Finges, souches pins (Pinus); 14.VIII.1980; leg.
C. Besuchet — (5).
NW-3
EIST OF IDENTIFIED SPECIES
OPPIIDAE Grandjean, 1951
Berniniella conjuncta (Strentzke, 1951) comb. et stat. n.
Locality: NW-2.
Distribution: Germany: first record for Switzerland.
Berniniella hauseri (Mahunka, 1974) comb. n.
Localities: GR-6; TI-11.
Distribution: Europe: first record for Switzerland.
ORIBATIDS FROM SWITZERLAND III 5]
Dissorhina signata (Schwalbe, 1989) comb. n.
Locality: GR-9.
Distribution: Germany (known from the type locality only); first record for
Switzerland.
Lauroppia hauseri sp. n.
Localities: TI-5; TI-11.
Lauroppia maritima (Willmann, 1929) comb. n.
Localities: SO-5; TI-9.
Distribution: Palearctic Region; first record for Switzerland.
Lauroppia obscura sp. n.
Localities: GR-8; VS-4; VS-8.
Moritzoppia incisa sp. n.
Localiy: NW-3.
Moritzoppia keilbachi (Moritz, 1969)
Locality: SZ-3.
Distribution: Europe; first record for Switzerland.
Oppiella besucheti sp. n.
Locality: SZ-3.
Oppiella propinqua sp. n.
Locality: LU-1.
Oxyoppioides decipiens (Paoli, 1908)
Locality: GE-4.
Distribution: Central and Southern Europe, Caucasus; first record for Switzerland.
Paramedioppia helvetica gen. n., sp. n.
Locality: TI-9.
Subiasella quadrimaculata (Evans, 1952)
Locality: GR-10.
Distribution: Europe; first record for Switzerland.
QUADROPPIIDAE Balogh, 1983
Quadroppia longisetosa Minguez, Ruiz & Subias, 1985
Locality: NW-1.
Distribution: Southern Europe; first record for Switzerland.
Quadroppia michaeli Mahunka, 1977
Locality: not recorded for Switzerland.
Distribution: Greece and Spain.
Quadroppia omodeoi Mahunka & Paoletti, 1984
Locality: not recorded for Switzerland.
Distribution: Italy.
Quadroppia cf. paolii Woas, 1986
Locality: GR-10.
Distribution: Palearctic Region; first record for Switzerland.
DESCRIPTION AND REDESCRIPTION OF SOME OF THE OPPIOID SPECIES
The present paper records 15 species belonging to the superfamily Oppioidea
Grandjean, 1951 found in the territory of Switzerland (13 species of the family Oppii-
dae and 4 species of the family Quadroppiidae). In exceptional cases we also discuss
species unrecorded in Switzerland so far, when it is deemed important to elucidate
certain specific taxonomic questions. Among the species 6 are new for science and |
represents a new genus (Paramedioppia). In addition some extremely rare species are
touched upon. Morphological complementations, corrections, new combinations, the
publication of synonyms and the reinterpretation of some species are also given.
52 SANDOR MAHUNKA & LUISE MAHUNKA-PAPP
Berniniella conjuncta (Strenzke, 1951) comb. et stat. n. Figs 1-3
Oppia sigma conjuncta Strenzke, 1951: 723, fig. 5.
Oppiella sigma (Strentzke, 1951): sensu Woas, 1986: 182, Abb. 88-89.
Material examined: Switzerland: NW-2.
Remarks: The recently examined soil samples harboured, in many cases,
the very same species which we could readily identify with the description of
Strenzke’s Oppia sigma conjuncta published in 1951, and with Woas's redescription
Fics 1-3
Berniniella conjuncta (Strenzke, 1951) — 1: body in dorsal view, 2: body in ventral view, 3:
podosoma in lateral view.
ORIBATIDS FROM SWITZERLAND III 53
of Oppia sigma published in 1986. Without doubt Strenzke's description is inadequate
since the description of the ventral side is missing. However, on the basis of the Swiss
specimens and the redescription of Woas now we may safely conclude that O.
conjuncta is an independent, valid species. Furthermore, we should like to point out
that Woas made a mistake when preparing his redescription, since his specimens
belong to O. conjuncta and not to O. sigma.
The drawings were made from the specimens originating from Switzerland
which readily correspond with the figures of both Strenzke and Woas. For this reason
a complete redescription would be out of place here. Nevertheless, we should like to
draw the attention to some important features.
Measurements: Length of body: 208-224 um, width of body: 102-
114 um.
Dorsal side (Fig. 1): Rostrum with wide median apex, lateral apices
much smaller. Costulae S-shaped, directed to the bothridia. One pair of median laths
directed outwards, not connecting with the costulae. Bothridium with posterior
tubercle, behind it comparatively large pustules are evident. Sensilli with short
bristles. Dorsosejugal suture protruding anteriorly, ten pairs of notogastral setae
nearly equal in length.
Lateral part of podosoma (Fig. 3): Well sclerotized, longitudinal
crests and pustulated or granulated fields are visible. Ratio of prodorsal setae: ro = ex
> in = le. Setae /c do not arise on pedotecta I, discidium well developed bearing
setae 4c.
Ventral side (Fig. 2): Sternal apodemes and borders weakly developed
or absent. Sejugal apodemes with longitudinal, arched lines in opposite position.
Posterior margin of epimeral borders 2 and 4 undulate or denticulate. Epimeres 1
framed laterally by longitudinal crests bearing setae /c anteriorly. Anogenital setal
formula: 4 - 1 - 2 - 3. All setae in the epimeral and anogenital region simple, setiform,
without conspicuous cilia.
Berniniella hauseri (Mahunka, 1974) comb. n. Figs 4-5
Oppia hauseri Mahunka, 1974: 585, Abb. 34-36.
Oppiella rafalskii Optotna & Rajski, 1983: 543, figs 1-8. Syn. n.
Material examined: Switzerland: GR-6; T-11.
Remarks: The recently collected specimens are quite identical with those
described from Greece (Mahunka, 1974). However, the ventral side was not
described, so the relegation of the species was only provisional owing to lack of
information regarding the number of genital setae and some other ventral features.
After having again examined the type species we found, as it was expected from the
dorsal side, that O. hauseri has 4 pairs of genital setae corresponding well in all other
features with those of specimens collected recently in Switzerland. One of these was
selected for the drawings (Figs 4-5).
Taxonomic position: After examining the related species, it is
certain that Berniniella serratirostris (Golosova, 1970) comb. n. and Berniniella
54 SANDOR MAHUNKA & LUISE MAHUNKA-PAPP
RE
\ )
Fics 4-5
Berniniella hauseri (Mahunka, 1974) — 4: body ın dorsal view, 5: body in ventral view.
rafalskii (Optotna & Rajski, 1983) comb. n. are its closest allies. Peculiarly enough,
we could not find any differentiating character between the latter and B. hauseri, thus,
we consider them synonyms; so the latter is a junior synonym of B. hauseri. The
authors’ description - besides the obvious mistakes - 1s wholly identical with the
features of the Swiss specimens. The only difference we could ascertain stems from
the drawing technique, due to some simplified method used by the authors. Thus, the
median part of the dorsosejugal region, the costula on the basal part of the prodorsum
and also the median thickening are figured to be more complicated than they in fact
are. A similar problem is found with the drawing of the sejugal borders and the
coxisternal region. The position of setae 2b, 2c and 3c is obviously erroneously
depicted. Optotna & Rajski (1983) thoroughly evaluated the relationship of Oppia
serratirostris with Oppiella rafalskii. These differences (especially the number of
branches of the sensilli, the shape of the costulae and the shape of the median rostral
apex) are convincing enough for us, too.
Dissorhina signata (Schwalbe, 1989) comb. n. Figs 6-9
Oppia signata Schwalbe, 1989: 99, Abb. 1.
Material examined: Switzerland: GR-9.
ORIBATIDS FROM SWITZERLAND III
Nn
Un
Fics 6-7
Dissorhina signata (Schwalbe, 1989) — 6: body in dorsal view, 7: body in ventral view.
Remarks: On the basis of the sensillar shape and its unilateral ciliation,
the species of the genus Dissorhina Hull, 1916 may grouped into two. The “ornata
species group” has bacilliform sensilli, whose distal end is spiculate or quite smooth,
while the “fricarinatoides species group” has sensilli with very long unilaterally
arranged branches. The studied species belongs to the second group.
Redescription: The specimens of this species collected in Switzerland
have been stored for some time in the Geneva collection. These specimens are easily
identifiable with the description and the given figures. Some important features which
deserve special attention are described below.
Measurements. — Length of body: 197-208 um, width of body: 103-
134 um.
Dorsal side (Fig. 6): Rostrum tripartite, rostral setae arising exactly on
the triangular, well separated median apex. Costulae casket-shaped, diverging me-
dially and converging distally. Lamellar setae arising on small separated parts. Bothri-
dium with a posterior tubercle, behind it a well developed, separated, round tubercle.
Sensilli with 6-7 long, branches unilaterally. Dorsosejugal suture well protruding
anteriorly, with a straight median part. Ten pairs of notogastral setae nearly equal in
length. Setae c, not shorter than the others.
56 SANDOR MAHUNKA & LUISE MAHUNKA-PAPP
Fics 8-9
Dissorhina signata (Schwalbe, 1989) — 8: podosoma in lateral view, 9: leg I.
Lateral part of podosoma (Fig. 8): Well sclerotized, longitudinal
crests and pustulated or granulated fields are visible. Ratio of prodorsal setae: ro = ex
>in = le. Setae /c do not arise on pedotecta I, discidium well developed and bearing
setae 4c.
Ventral side (Fig. 7): Sternal apodemes absent between bo. 2 and bo.
sej. Sejugal apodemes with longitudinal, arched lines. Posterior margin of epimeral
borders 4 undulate or denticulate. Epimeres 1 framed laterally by longitudinal crests
bearing setae /c. Anogenital setal formula: 5 - 1 - 2 - 3. All setae in the epimeral and
anogenital region simple, setiform, without conspicuous cilia.
Legs: Form and chaetotaxy of leg I as shown in Fig. 9.
Taxonomic position: The species D. signata stands very near to D.
tricarinatoides (Dubinina, in Dubinina er al. 1966) (see also Mahunka, 1996b). It may
be distinguished by the shape of the prodorsal costulae with the auxiliary comple-
mentary ribs, which are present in D. tricarinatoides and absent in D. signata, and the
position of setae ad, which arise nearer to the anal plates than the lateral margin of
ventral plate in D. signata (much farther, near to the lateral margin in D. trticari-
natoides).
ORIBATIDS FROM SWITZERLAND III
57
Fics 10-12. Lauroppia hauseri sp. n. — 10: body in dorsal view, 11: body in ventral view,
12: podosoma in lateral view.
Lauroppia hauseri sp. n. Figs 10-12
Material examined: Switzerland: Holotype: Tessin: TI-11, 1 paratype from
the same sample; 6 paratypes: Tessin: TI-5. Holotype and 4 paratypes: MHNG!, 3 paratypes
(1614-PO-98): HNHM?.
' MHNG: deposited in the Muséum d’histoire naturelle, Geneva.
2 HNHM: deposited in the Hungarian Natural History Museum, Budapest, with identification
number of the specimens in the Collection of Arachnida.
58 SANDOR MAHUNKA & LUISE MAHUNKA-PAPP
Diagnosis: Rostrum elongated, rostral apex roundish. Short, basally
dilated, guttiform costulae present Sensilli pectinate. Median part of the anterior
notogastral margin straight. Ten pair of notogastral setae, setae c, long. Six pairs of
genital setae, anogenital region normal.
Measurements: Length of body: 369-443 um, width of body: 188-
214 um.
Prodorsum: Rostral part of prodorsum conspicuously elongated, rostral
apex conical, rounded medially. Prodorsal surface with short basal transcostulae pro-
truding from the guttiform basal tubercles. Bothridia well developed, with conspi-
cuously large basal tubercles. Rostral setae arising from the rostral surface, lamellar
setae located very far from the rostrals, near to the interlamellar setae. Lamellar setae
fine, setiform, arising at the distal end of the costulae, interlamellar setae much
thicker, ciliate, straight, arising from the guttiform tubercles. Sensilli pectinate, direc-
ted forwards, each bearing 7 branches unilaterally.
Lateral part of podosoma: Sejugal region covered by a thick
cerotegument layer. Surface weakly granulated, with larger pustules between the
acetabula. Acetabula IV placed above acetabula I-III. Setae /c arising near to small
pedotecta I (Fig. 12).
Notogaster: Anteromedian margin of notogaster straight, not penetrating
between the bothridia. A weak crista present. Ten pairs of notogastral setae present,
setae c, long, longer than the others. Setae /a arising in front of Im (Fig. 10). Setae la
and h; longer than the remaining ones, setae p,, p,, and p3 shortest of all.
Ventral side (Fig. 11): Apodemes and borders weakly developed,
epimeral surface ornamented by a weak polygonal design. Posterior border of epi-
meral region undulate or denticulate. Epimeral setae partly ciliate or bifurcate. Lateral
setae (3c, 4c) longer than the inner ones. Anogenital setal formula: 6 - 1 - 2 - 3. All
setae in anogenital region simple and short.
Remarks: On the basis of the prodorsal structure and the basal position of
the lamellar and interlamellar setae the new species belongs to the “fallax-group” (see
Willmann, 1931). The new species is distinguished from its congeners by the ratio
and form of the prodorsal setae and the very short / and p setae.
Derivatio nominis: We dedicate the new species to our friend, Dr. B.
Hauser, the initiator of the “Oribatida Helvetica project”.
Lauroppia maritima (Willmann, 1929) comb. n. Fig. 13-15
Oppia maritima Willmann, 1929: 45, Abb. 4.
Oppia maritima: Strenzke, 1951: 720, fig. 2
Material examined: Switzerland: SO-5; TI-9.
Remarks: The species was described by Willmann (1929), and Strenzke
(1951) redescribed it. Both these descriptions are inadequate and some very important
features are missing. Therefore, we give herewith some drawings and a short des-
cription.
ORIBATIDS FROM SWITZERLAND III 59
Fics 13-15
Lauroppia maritima (Willmann, 1929) — 13: prodorsum in dorsal view, 14: epimeral region,
15: podosoma in lateral view.
Prodorsum: Rostral apex sharply pointed, the incisure wide, lateral
apices much smaller than the median one (Fig. 13). Exobothridial setae bifurcate.
Exobothridial region well sclerotized (Fig. 15), surface around the bothridia and
acetabulae I-IV pustulate or granulate.
Notogaster: Protruding anteriorly, however, the median part of the
dorsosejugal line is straight.
Ventral side: Posterior border of the coxisternal region with pustulate
margin, pustules are in a transversal hollow on both sides (Fig. 14). Setae /c arising
far from pedotecta I. Setae 3c bifurcate. Anogenital setal formula: 5 - 1 - 2 - 3.
Anterior pair of genital setae much longer than the others.
60 SANDOR MAHUNKA & LUISE MAHUNKA-PAPP
Lauroppia obscura sp. n. Figs 16-20
Material examined: Switzerland: Holotype: Valais: VS-8, 6 paratypes from
the same sample; 4 paratypes: Valais: VS-4; 6 paratypes: Graubünden: GR-8. Holotype and 10
paratypes: MHNG, 6 paratypes (1615-PO-98): HNHM.
Diagnosis: Rostrum divided by two incisions, median apex triangular.
Prodorsum with complicate costulae. Sensilli clavate, unilaterally spinose. Notogaster
straight anteriorly. Ten pairs of notogastral setae, c, long. 5 pairs of genital setae,
anogenital region normal.
Measurements: Length of body: 280-305 um, width of body:
148-172 um.
Prodorsum: Rostral apex (Fig. 17) divided by two small incisions,
median apex conspicuously wide, sometimes triangular, or blunt at tip, sometimes its
margin undulate. Lateral apices with simple, sharp points. Among the costulae one
pair arched and comparatively well developed, they do not reach the bothridium or
the bases of the lamellar setae. A pair of S-shaped basal costulae also present
reaching to the inner margin of the longitudinal ones (Fig. 20). Lateral part of
prodorsum well sclerotized, with some tubercles and pustules, one pair of which bears
the exobothridial setae. In the dorsosejugal region one pair of indistinct porose areas
also visible (Fig. 19). Bothridia large with strong posterior apophyse. Rostral setae
arising dorsally, comparatively near to each other. Lamellar setae shorter than the
other prodorsal setae, arising on small tubercles independently of the longitudinal
costula. Sensilli elongate, their head asymmetrically clavate, with short spines on their
outer margin.
Lateral part of podosoma: Exobothridial region and fields
between acetabula I - IV granulated by comparatively large granules or pustules.
Some well-sclerotised, long, longitudinal crests also present. A pair of porose areas in
the sejugal region. Positions of acetabula III and IV nearly the same as those of ace-
tabula I and II (Fig. 19). Pedotecta I small, setae /c arising far from its basal part.
Discidium well developed, setae 4c arising far from its lateral margin, on the epimeral
surface.
Notogaster: Elongated, dorsosejugal part straight anteriorly. Crista
clearly seen. Ten pairs of notogastral setae present, setae c, conspicuously long,
setiform, all others somewhat shorter. Setae /m arising in front of la (Fig. 16). Setae
pP], P and pz only slightly shorter than the others.
Ventral side (Fig. 18): Epimeral region hardly sclerotised. Apodemes
partly absent only borders visible between the borders 2 and sejugal borders or sejugal
borders and borders 4. Lateral borders on epimeres | arched bearing setae /c. Sejugal
borders broad, with a pair of hollows medially, with similar, but smaller ones, also
present on bo 2. All epimeres with a polygonal pattern. Setae 4c arising on the
epimeral surface. Epimeral setae long, simple. Anogenital setal formula: 5 - 1 - 2 - 3.
Among the genital setae the two anterior pairs conspicuously long, directed forwards.
Position and shape of the aggenital, anal setae and lyrifissures iad normal.
Legs: Solenidia of tibia IV very long and curved .
ORIBATIDS FROM SWITZERLAND III 61
Fics 16-20
Lauroppia obscura sp. n. — 16: body in dorsal view, 17: rostrum, 18: body in ventral view,
19: podosoma in lateral view, 20: basal part of prodorsum.
62 SANDOR MAHUNKA & LUISE MAHUNKA-PAPP
Remarks: On the basis of the form and structure of the prodorsum the new
species belongs to the L. maritima group. Although closely related to L. maritima, the
head of the sensillus of the new species is much longer and the spines on the sensillar
head are very short (long in L. maritima).
Derivatio nominis: Named after the complicated prodorsal design
and rostral apex.
Moritzoppia incisa sp. n. Figs 21-23
Materialexamined: Switzerland: Holotype: Nidwald: NW-3, 26 paratypes
from the same sample. Holotype and 16 paratypes: MHNG, 10 paratypes (1613-PO-98):
NHMH.
Diagnosis: Rostrum roundish. Prodorsum with complicate costulae.
Sensilli clavate, unilaterally pectinate. Anterior margin of notogaster protruding into
the interbothridial region and interrupted medially. Cristae present. Ten pairs of
notogastral setae, setae c, long. Four pairs of genital setae, anogenital region normal.
Measurements: Length of body: 267-276 um, width of body: 152-
155 um.
Prodorsum: Rostrum slightly elongated, with nasiform median apex.
Lamellar costulae weakly sclerotized, lamellar apices usually separated, their anterior
part carrying some pustules. A pair of elongated, guttiform tubercles in the inter-
bothridial region bearing interlamellar setae at their outer margin. A pair of round
tubercles present in the middle part of the costulae, they are opposed on both sides
(Fig. 21). Some weak sigillae visible laterally. Bothridia well developed, with large
basal tubercles surrounded with a granulated field. Rostral setae arising on the dorsal
surface, ciliate. Lamellar setae arising on the lamellar apices. The ratio of the pro-
dorsal setae: ro > in = ex > le. Sensilli with well developed heads, bearing 9-10
branches unilaterally.
Lateral part of podosoma: Well sclerotized, nearly the whole
surface well granulated and/or pustulate. Large pustules present also above the ace-
tabula of legs HI and IV (Fig. 23). Some longitudinal crests also present. Ventral
border especially undulate and incised, acetabula I-IV arranged nearly in a longitu-
dinal line. No porose area visible behind the bothridia.
Notogaster: Dorsosejugal part of the notogaster protruding anteriorly in
the interbothridial region. Crista present laterally. Dorsosejugal margin interrupted
medially and framed by a pair of short crests directed posteriorly, sometimes some
spots are visible in this slit. Ten pairs of notogastral setae present, setae c, slightly
longer than the others, la, Im and /p equal in length, setae h and setae p only slightly
shorter.
Ventral side (Fig. 22): Well sclerotized, all apodemes and borders well
developed. Lateral margin of epimeres | framed by longitudinal borders, but setae /c
arising on pedotecta I. The surface of epimeres I pustulate, posterior borders also
conspicuously covered by granules or pustules. Sejugal borders with some pairs of
longitudinal crests or lines. Posterior margin of borders IV strongly undulate. Setae 4b
arising at the posterolateral corner. Surface around discidium also pustulate. Setae 4c
ORIBATIDS FROM SWITZERLAND III 63
Fics 21-23
Moritzoppia incisa sp. n. — 21: body in dorsal view, 22: body in ventral view, 23: podosoma in
lateral view.
64 SANDOR MAHUNKA & LUISE MAHUNKA-PAPP
stand further from the discidia. All epimeral setae simple and smooth. Anogenital
setal formula: 4 - 1 - 2 - 3. All setae simple. Lyrifissures iad short, located in paraanal
position.
Remarks: On the basis of the structure of the dorsosejugal margin of the
notogaster, the new species is distinguished from all congeners.
Derivatio nominis: Named after the incised anterior margin of the
notogaster.
Moritzoppia keilbachi (Moritz, 1969)
Oppia keilbachi Moritz, 1969: 37, Abb. 1-3.
Moritziella keilbachi: Balogh, 1983: 26, 9: 9p-v.
Moritzoppia keilbachi: Subias & Rodriguez, 1988: 122.
Material examined: Switzerland: SZ-3.
Remarks: The species was described by Moritz (1969) on the basis of
females. A male specimen has been found in the Swiss material. The habitus of the
male is more slender and it is also smaller (210um x 92 um) than the female. In all
other features it corresponds well with the original description and the figures. Espe-
cially important is the granulate surface behind the bothridium and on the discidium
and the shape of the sensillus. The sensillus of the studied individual is blunt at tip, its
head bearing simple, short cilia, arranged in 2-3 rows. Features of the ventral region
also accord well with the original description, e.g. the posterior borders of the
coxisternal region have an undulate margin.
Oppiella besucheti sp. n. Figs 24-27
Material examined: Switzerland: Holotype: Schwyz: SZ-3, 5 paratypes from
the same sample. Holotype and 3 paraypes: MHNG, 2 paratypes (1611-PO-98): NHMH.
Diagnosis: Rostral apex divided by two incisions, median apex
triangular. Prodorsum with costulae and other structures. Sensilli narrow, fusiform
with lateral branches. Anterior margin of notogaster straight, with lateral protube-
rances. Ten pairs of notogastral setae, setae c, long. Five pairs of genital setae,
anogenital region normal.
Measurements: Length of body: 250-259 um, width of body: 99-
105 um.
Prodorsum: Rostrum divided by two small, but wide, U-shaped
incisions, median apex triangular, also small (Fig. 24). Median costulae Y-shaped,
weakly developed. Behind them, in the interbothridial position, one pair of elongated,
nearly guttiform tubercles. Along the lamellar costulae, at their outer side, a pair of
opposed smaller tubercles present on each side. Bothridia with posterior tubercles.
The whole surface of prodorsum distinctly granulated or pustulated, also in the
lamellar region (Fig. 27). All four pairs of prodorsal setae long, interlamellar ones
thicker and more heavily ciliated than the others. Ratio among them: in > ro = ex >
le. Sensilli very long, slightly dilated medially, bearing 5-6 lateral branches of
different lengths on their distal parts.
ORIBATIDS FROM SWITZERLAND III 65
Fics 24-27
Oppiella besucheti sp. n. — 24: body in dorsal view, 25: body in ventral view, 26: podosoma in
lateral view, 27: basal part of prodorsum.
66 SANDOR MAHUNKA & LUISE MAHUNKA-PAPP
Lateral part of podosoma: Well sclerotized, nearly the whole
surface well granulated and/or pustulated. Some longitudinal crests also present.
Ventral border especially waved and incised, acetabula I-IV arranged nearly in one
longitudinal line (Fig. 26). A pair of porose areae behind the bothridia clearly seen.
Notogaster (Fig. 24): With one pair of granulate humeral apophyses.
Dorsosejugal suture straight. Ten pairs of notogastral setae present, setae c, shorter
than Ja, Im and Ip also shorter than c,, but nearly equal in length with the other setae,
excepting p, and p3. Setae h, characteristically directed outwards. Setae c, with cilia,
all the other setae smooth.
Ventral side (Fig. 25): Well sclerotized, all apodemes and borders -
excepting a part of the sternal ones - between the 2nd and sejugal apodemes well deve-
loped. Lateral margin of epimeres | framed by longitudinal borders bearing setae /c. A
pair of inner costulae, parallel with discidium also present on epimeres 4 continued as
undulate transversal parts. Setae 4b arising on their posterolateral corner. Among the
epimeral setae, setae /c, 3c, but primarily and firstly setae 4c, with long cilia. All other
epimeral setae simple and smooth. Anogenital setal formula: 5 - 1 - 2 - 3. All setae
simple. Lyrifissures iad long, placed in paraanal position.
Legs: Setae of trochanters of legs II well arched, with conspicuously long
cilia. Setae p of all tarsi simple, setiform. Solenidium @ of leg IV, very long, arched,
longer than the joint.
Remarks: See after the following species.
Derivatio nominis: I dedicate the new species to Dr. C. Besuchet
(Muséum d'histoire naturelle, Geneva), the collector of the specimens and most of
those soil samples from Switzerland which we studied recently in Geneva.
Oppiella propinqua sp. n. Figs 28-32
Material examined: Switzerland: Holotype: Luzern: LU-1 15 paratypes from
the same sample. Holotype and 10 paratypes: MHNG, 5 paratypes (1612-PO-98): NHMH.
Diagnosis: Rostrum nasiform. Lamellar costulae directed to the
bothridia, one pair of basal protuberances present. Sensilli slightly fusiform, narrow,
bearing short spines. Anterior margin of the notogaster straight, with well developed
lateral protuberances. Ten pairs of notogastral setae, setae c, long. Anogenital region
typical for the genus. Five pairs of genital setae.
Measurements: Length of body: 263-270 um, width of body: 140-
145 um.
Prodorsum: Rostrum slightly elongated, with a nasiform median apex.
Lamellar costula well sclerotized, covered by pustules on their anterior part. A much
finer transversal costula between them (sometimes it is hardly observable) (Fig. 30).
A pair of elongated, guttiform tubercles in the interbothridial region bearing inter-
lamellar setae on their outer margin and a pair of short laths at an angle between
costulae and bothridia. Some weak sigillae visible laterally. Bothridia well developed,
with a basal tubercle with a granulated field around it. Rostral setae characteristically
arched inwards, ciliate. Lamellar setae arising on the distal half of the costulae. The
ORIBATIDS FROM SWITZERLAND III 67
Fics 28-29
Oppiella propinqua sp. n. — 28: body in dorsal view, 29: body in ventral view.
ratio of the prodorsal setae: ro > in = ex > le. Sensilli very long, their heads slightly
dilated medially bearing some (5-7) very short spines.
Lateral part of podosoma (Fig. 32): Well sclerotized, nearly the
whole surface well granulated and/or pustulated. Some longitudinal crests also
present. Ventral border, especially, undulate and incised acetabula I-IV arranged in
one longitudinal line. A pair of indistinct porose areas behind the bothridia.
Notogaster (Fig. 28): With one pair of large humeral apophyses.
Dorsosejugal suture straight medially, slightly arched laterally, near the humeral
tubercles. Ten pairs of notogastral setae present, setae c, and /a equal in length, /m
and /p much shorter than the preceding ones. Setae h, characteristically directed
outwards, no essential difference among setae h and setae p,. Setae c, with short cilia,
all other setae smooth.
Ventral side (Fig. 29): Well sclerotized, all apodemes and borders -
excepting a part of the sternal ones - between the 2nd and sejugal apodemes well
developed. Lateral margin of epimeres | framed by longitudinal borders bearing
68 SANDOR MAHUNKA & LUISE MAHUNKA-PAPP
Fics 30-32
Oppiella propinqua sp. n. — 30: basal part of prodorsum, 31: epimeral region, 32: podosoma in
lateral view.
setae /c. A pair of inner costulae, parallel with the discidium also present on epi-
meres 4 continued as undulate transversal parts (Fig. 31). Setae 4b arising on their
posterolateral corner. All epimeral setae simple and smooth. Anogenital setal formula:
5 - | - 2-3. All setae simple. Lyrifissures iad long, located in paraanal position.
Legs: Setae p of legs II - IV thick, clearly spiniform. Solenidium of tibia IV
not longer than the joint.
Remarks: Oppiella nova (Oudemans, 1902) is one of the most variable
species of the genus Oppiella Jacot, 1937, however, it was designated as the type of
the genus. Most authors consider this species as cosmopolitan, but nobody has made a
comparative study on world material. Our opinion is, that this species comprises a
number of closely related species. This opinion is strengthened in that some authors
consider Oppiella uliginosa (Willmann, 1919) as a synonym of O. nova (e.g. Subias
& Balogh, 1989). On the other hand Woas (1986) in his redescription of O. uliginosa
quite obviously represents another species. The description and the drawings (Woas,
1986: 208, Abb. 102-103) surely do not refer to O. nova.
ORIBATIDS FROM SWITZERLAND III 69
The two preceding new species stand very near to the type species of this
genus, and e.g. Oppiella primorica (Golosova, 1970), which also belongs to this
group. I also regard this genus as rather heterogeneous needing further study to
demarcate its generic limits. Obviously, on the basis of the O. nova type, the diag-
nosis of the genus Oppiella should be complemented and restricted so that only those
species which display the following characteristics should be included: Dorsosejugal
margin with lateral apophyses. Among the epimeres, epimeres | framed laterally with
a longitudinal border bearing setae /c, epimeres 4 framed posteriorly by an undulate
(denticulate) lath, continuing anteriorly as a longitudinal inner costula. Other features
should be investigated in the future.
Among the new species, on the basis of the undivided rostrum, Oppiella pro-
pinqua sp. n. stands closer to Oppiella nova. It is distinguishable from the type spe-
cies by the form of the sensillus, which is longer in the new species. The other
differential characters are: form of the prodorsal costula, the form and ratio of pro-
dorsal setae, length and ciliation of epimeral setae (/c, 3c and 4c). The other species
(Oppiella besucheti sp. n.) stands nearest to Oppiella primorica (Golosova, 1970) on
the basis of the divided rostral apex (see also Mahunka, 1979). The new species is
distinguishable from Oppiella primorica by the much larger median prodorsal apex
and the thicker and ciliated interlamellar setae and the much wider basal guttiform
tubercles in the interbothridial region in the new species.
Derivatio nominis: The new species belongs to the Oppiella nova
species group.
Oxyoppioides decipiens (Paoli, 1908) Figs 33-37
Oppia decipiens Paoli, 1908: 69, figs 29, 48.
Oppia decipiens: Pérez-Inigo, 1971: 297, fig. 30.
Oxyoppioides decipiens: Subias & Minguez, 1985: 182.
Material examined: Switzerland: GE-4.
Remarks: The species has been collected several times in Central Europe,
although it is more common in the Mediterraenean area. Several authors (e.g. Pérez-
Inigo, 1965; Subias & Minguez, 1985) discussed it but no detailed redescription has
been made. Furthermore,there are no available figures illustrating its leg structure.
The following remarks complement our knowledge of this species.
Prodorsum: Rostrum U-shaped, excavated with two small, lateral teeth
(Fig. 33). Costulae weak, surface densely granulate. Interbothridial region with
numerous indistinct sigilla.
Pate ral part of pod os om a. (Eis. 35): Weakly, sclerotized:
exobothridial region with fine granulation, with only poorly developed, longitudinal
laths. Exobothridial setae short.
Notogaster: Humeral processes in the dorsosejugal region, hitherto
characteristic for the genus Oxyoppia, entirely missing. Anterior margin of notogaster
arcuate, closed in the middle.
Ventral side (Fig. 34): Apodemes very poorly developed. Epimeral
borders hardly discernible in places. From among the sejugal and apodemes 4 and
70 SANDOR MAHUNKA & LUISE MAHUNKA-PAPP
Fics 33-35
Oxyoppioides decipiens (Paoli, 1908) — 33: body in dorsal view, 34: body in ventral view, 35:
podosoma in lateral view.
borders the sternal apodeme and border are entirely missing. Surface of discidium
granulate. Setae /c arising on pedotecta 1. Epimeral setae ciliate, setae 4a and 4b
located strikingly close to each other. Anogenital formula 5-1-2-3. Aggenital setae
arising one behind the other. Lyrifissure iad in inverse apoanal position, far removed
from anal opening.
ORIBATIDS FROM SWITZERLAND III il
Legs: Normal, oppioid type, only the tibia of leg I slightly thickened and
the distal part of femur triangular (Fig. 36), solenidium of tibia of leg IV (Fig. 37)
conspicuously short, and all setae characteristically long.
Taxonomic position: Subias & Balogh (1989) placed the genus in
relationship with Oxyoppia. We do not agree with this opinion, although the genus is
soundly based. The key published by them is unsuitable for its identification. The
position of lyrifissure fad, said to be paraanal, is also in error in the key of Balogh &
Balogh (1992).
FIGS 36-37
Oxyopioides decipiens (Paoli, 1908) — 36: leg I, 37: femur, genu and tibia of leg IV.
Paramedioppia gen. n.
Diagnosis: Family Oppidae. Rostrum divided by two incisions. Weak
costulae with a transcostula and one pair of interbothridial “tubercles” present on
prodorsum. Sensilli setiform, with some irregular spines. Dorsosejugal part of noto-
gaster gradually arched, not penetrating into the interbothridial region. Crista present,
setae c, arising medially. Ten pairs of notogastral setae present. Among the epimeral
Tp. SANDOR MAHUNKA & LUISE MAHUNKA-PAPP
setae /c arising on the epimeral surface, epimeral borders and apodeme normally
developed. Anogenital setal formula 6 - 1 - 2 - 3. Lyrifissures iad in paraanal position.
Gnathosoma, chelicerae and palps normal. Legs very long, joints of leg IV especially
elongated.
Type species: Paramedioppia helvetica sp. n.
Remarks: The new taxon seems to be a mixture of two genera: Oppiella
Jacot, 1937 and Medioppia Subias & Minguez, 1985. The latter is a highly hetero-
geneous genus, as is Oppiella. The type species of the genus Medioppia, O. media
Mihelcic, 1956 (Pérez-Iñigo, 1965) does not have costulae but has interbothridial
tubercles. On the other hand, it has 5 pairs of genital setae, and its rostrum is
undivided. Thus, we could not relegate the new species into any of the known genera.
Derivatio nominis: Similar to the genus Medioppia.
Paramedioppia helvetica sp. n. Figs 38-42
Materialexamined: Switzerland: Holotype: Tessin: TI-9, 1 paratype from
same sample. Holotype: MHNG, paratype (1616-PO-98): NHMH.
Diagnosis: Rostrum divided, rostral apex truncate. Prodorsum weakly
sclerotized, short longitudinal and transversal costulae present. Sensilli setiform with
irregular spines. Anterior margin of notogaster convex. Cristae present. Ten pairs of
notogastral setae, setae c, long. Anogenital setae normal, six pairs of genital setae.
Measurements: Length of body: 762-823 um, width of body: 369-
403 um.
Prodorsum: Elongated. Rostral part rounded, but flanked by two short
incisions. Median apex truncate, lateral apices rounded. Prodorsal surface weakly
sclerotized, but short longitudinal costulae with distinct corners, a very weak trans-
costula and a pair of interbothridial, arched tubercles present. Some indistinct sigilla
also observable. Bothridia without posterior tubercles. Rostral setae arising on the
prodorsal surface, lamellar setae located behind the costular corners (Fig. 38). Ratio
of prodorsal setae: : in = ro > le > ex. Sensilli well characteristic for the species,
bent several times, setiform, with 4-5 spines, without capitula, their distal ends simple
or bifurcate.
Lateral part of podosoma: Pedotecta I comparatively well
developed. Exobothridial surface not granulate, some granules observable betweeen
the acetabula II - HI. This part well framed by a lath, above the acetabula (Fig. 40).
Notogaster: Conspicuously elongated. Crista weak, but distinct also a
pair of short and weak median lines present at the dorsosejugal margin of the noto-
gaster (Figs. 38, 40). Ten pairs of notogastral setae of equal length present, all ciliate.
Ventral side (Fig. 39): Apodemes and borders weakly developed.
Marginal, longitudinal lath absent on epimera 1. In anteromedian sternal apodemes a
ring-like feature present on the sternal apodemes, between setae /a. All epimeral
setae simple, thin, setiform. Epimeral surface ornamented by polygonal pattern. Six
pairs of genital setae arising in two rows (4 pairs in median, 2 pairs in lateral
position). Adanal setae ciliate, like the notogastral ones, other setae in the anogenital
region simple.
ORIBATIDS FROM SWITZERLAND III 73
Fics 38-40
Paramedioppia helvetica gen. n., sp. n. — 38: body in dorsal view, 39: body in ventral view,
40: podosoma in lateral view.
Legs: All joints long, not widened, setae conspicuously ciliate/spinose. Leg
setal formulae are normal for the family:
I: 1-5-2+1-4+2-20+2 - 1 (Fig. 41)
IV:1-2-2-3+1-10-1 (Fig. 42)
74 SANDOR MAHUNKA & LUISE MAHUNKA-PAPP
Fics 41-42
Paramedioppia helvetica gen. n., sp. n. 41: leg I, 42: leg IV.
Remarks: The main features are mentioned in the generic diagnosis, but it
is the form of the sensillus which well characterises the new species.
Derivatio nominis: Named after the country the species originates
from.
Subiasella (Lalmoppia) quadrimaculata (Evans, 1952)
n
Oppia quadrimaculata Evans, 1952: 37, fig. 2.
Subiasella (Lalmoppia) quadrimaculata: Subias & Rodriguez, 1986: 114, figs 1-3.
Materialexamined: Switzerland: GR-10.
Remarks: The species is known from several localities (from England to
Poland) of Europe. but it is not frequent anywhere. This is its first record from
Switzerland. The prodorsum of these specimens, as it is the case in the Hungarian
ones, has neither costula nor lamellar line.
ORIBATIDS FROM SWITZERLAND III 75
Quadroppiidae Balogh, 1983
There is a great confusion in the family Quadroppiidae partly due to the
publication of Minguez et al. (1985) and partly to Woas (1986) but specifically to that
of the latter author who synonymized species without the study of types. The core of
the problem is in the interpretation of the senior author's species (Quadroppia
michaeli Mahunka, 1977) described from Greece, and on the other hand, Woas' inter-
pretation of the type species, ©. quadricarinata (Michael, 18853) which was misun-
derstood by Paoli (1908). Accordingly, if the ©. michaeli species is not new, but
simply the misinterpreted ©. quadricarinata, then the name introduced by Woas as
Q. paolii nom. n. is redundant.
The situation is further aggravated in that the senior author described with
Paoletti (Mahunka & Paoletti, 1984) another new species belonging to this genus (Q.
omodeoi Mahunka & Paoletti, 1984), without taking the trouble to point out its
relationship with the ©. quadricarinata by then correctly understood. Our recent
study has revealed that Q. paolii is probably identical with the specimens from
Switzerland, and we further believe that it is not identical either with ©. quadri-
carinata sensu Mahunka, nor with the form interpreted by Paoli, nor Q. omodeoi.
There is a further problem in that Minguez er al. (1985) hold the view that Hammer’s
Q. monstruosa is the same of Paoli’s Q. quadricarinta.
To clarify the problem thorough study of types is needed along with authenti-
cally identified specimens. This time we could only examine the types of Q. michaeli
and Q. omodeoi, and those Greek specimens which were published in 1977 as
O. quadricarinata sensu Mahunka. We discuss O. longisetosa Minguez, Ruiz et
Subias, 1985 that was also found in Switzerland (Q. quadricarinata (Michael, 1885)
was also collected at several localities in Switzerland) but refrain from discussing the
other species here.
Quadroppia longisetosa Minguez, Ruiz & Subias, 1985 Figs 43-44
Quadroppia longisetosa Minguez et al., 1985: 104, figs 6-7.
Material examined: Switzerland: NW-1.
Remarks: This species is readily identifiable among the members of the
genus Quadroppia Jacot, 1939. The Swiss specimens correspond well with the des-
cription and the figures of the species. Some slight differences exist, like the weak
longitudinal lath in the interlamellar region parallel with the costulae, and the
notogastral setae are somewhat more rigid than depicted in the drawing. We provide
some figures obtained from specimens taken in Switzerland .
Quadroppia michaeli Mahunka, 1977 Fig. 45
Quadroppia michaeli Mahunka, 1977: 914, Abb. 12.
Quadroppia michaeli: Minguez et al., 1985: 114, figs 17-18.
Quadroppia michaeli: Mahunka, 1977 sensu Woas, 1986: 215.
> The original description appeared in 1885 and not in 1887 as mentioned by Woas (1986).
76 SANDOR MAHUNKA & LUISE MAHUNKA-PAPP
SSA
D Sal
A
| Tape
Fics 43-44
Quadroppia longisetosa Minguez, Ruiz & Subias, 1985 — 43: body in dorsal view, 44: body in
ventral view.
Remarks: In spite of Woas’ opinion, this is a valid, independent species. It
cannot be brought into close relationship with any of the so far described species. Its
characteristics are:
Prodorsum: Oval crest in the rostral region undivided and gradually
becoming narrower basally. On both sides an arcuate, well separated crest present.
Intercostular region with a short, transverse lath directly behind the distal end of
costulae.
Notogaster: The posteriorly running ribs emanating from the humeral
processes are not weaker than the ones beside them, length of ribs is the same.
Ventral side: Epimeral borders are shown in Fig. 45. The median
pattern directly in front of the genital opening is highly characteristic, it comprises
two touching parts, both are rounded.
Legs: Tarsus of leg II bears 2 solenidia.
Quadroppia omodeoi Mahunka & Paoletti, 1984 Figs 46-47
Quadroppia omodeoi Mahunka & Paoletti, 1984: 114, figs 1-2.
ORIBATIDS FROM SWITZERLAND III 77)
Remarks: All the known species are distant relatives only. Its charac-
teristics are:
Prodorsum: Oval crest in the rostral region is undivided, not strongly
narrowing basally. Intercostular region with a short, concave, transverse lath directly
behind the distal end of costulae. Behind this transverse lath 2 pairs of sigillae and
one pair of lateral laths are present, the latter are not connected in the middle. Sensilli
conspicuously long (Fig. 47), much longer than in the other species.
Notogaster: The posteriorly running ribs emanating from the humeral
processes are much longer than the lateral ones. Notogaster completely enframed by
arıb.
Ventral side: Fig. 46 depicts the epimeral borders.
Legs: Tarsus of leg II bears 2 solenidia.
Quadroppia cf. paolii Woas, 1986 Fig. 48-49
Quadroppia paolii Woas, 1986: 78, figs 30-32.
Material examined: Switzerland: GR-10.
Remarks: Without studying the type the relegation of this species is
impossible. It is highly probable that this species may be identical with Q. monstruosa
Hammer, 1979 sensu Minguez er al. (1985). Fundamental differences are obvious:
Prodorsum: Oval crest in the rostral region is either entirely divided
basally or much variegated. Intercostular region with a long, robust, arcuate lath
directly before the end of costulae. Just behind this 2 pairs of sigillae and one pair of
lateral laths, the latter frequently connected in the middle with a weak, transverse lath.
Sensillar capitula oval (Fig. 49).
Fics 45-49
Quadroppia michaeli Mahunka, 1977 — 45: epimeral region.
Quadroppia omodeoi Mahunka & Paoletti, 1983 — 46: epimeral region, 47: sensillus.
Quadroppia cf. paolii Woas, 1986 — 48: epimeral region, 49: sensillus.
78 SANDOR MAHUNKA & LUISE MAHUNKA-PAPP
Notogaster: The posteriorly running ribs emanating from the humeral
processes are shorter than the lateral ones.
Ventral side: as shown in Fig. 48.
Legs: Tarsus of Leg II bearing 2 solenidia.
ACKNOWLEDGEMENTS
First and foremost we should like to thank the collector of this interesting
material, Dr. C. Besuchet. Our hearthy thanks are due to Dr. V. Mahnert and Dr. B.
Hauser for the opportunity offered to study the material. We should also like to thank
Dr. C. Lienhard for the good advice extended on several taxonomic questions. For
reading the manuscript, translating some parts and linguistically revising others we
thank Dr. L. Zombori. For the conscientious corrections and the good advice incor-
porated in the text we extend our sincere thanks to Dr. M. Luxton (National Museum
of Wales, Cardiff).
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(1920): 552-554.
WILLMANN, C. 1929. Neue Oribatiden II. Zoologischer Anzeiger 80: 43-46.
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Andrias 5: 21-224.
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REVUE SUISSE DE ZOOLOGIE 107 (1): 81-95; mars 2000
Rudolphiella szidati sp. n. (Proteocephalidea: Monticelliidae,
Rudolphiellinae) parasite of Luciopimelodus pati (Valenciennes,
1840) (Pisces: Pimelodidae) from Argentina with new observations
on Rudolphiella lobosa (Riggenbach, 1895)
Alicia A. GIL DE PERTIERRA! & Alain de CHAMBRIER?
! Departamento de Ciencias Biolögicas, Lab. 52, Facultad de Ciencias Exactas y
Naturales, Pab. II, Ciudad Universitaria, U.B.A., 1428 - Buenos Aires, Argentina.
e-mail: helminto @bg.fcen.uba.ar
2 Departement des Invertébrés, Muséum d'histoire naturelle, PO Box 6434, CH -1211
Geneva 6, Switzerland.
e-mail: alain.dechambrier @ mhn. ville-ge.ch
Rudolphiella szidati sp. n. (Proteocephalidea: Monticelliidae, Rudol-
phiellinae) parasite of Luciopimelodus pati (Valenciennes, 1840) (Pisces:
Pimelodidae) from Argentina with new observations on Rudolphiella
lobosa (Riggenbach, 1895). - Rudolphiella szidati sp. n. (Cestoda, Monti-
celliidae, Rudolphiellinae) is described from the anterior intestine of the
pimelodid fish, Luciopimelodus pati caught in Parana river and de la Plata
river (Argentina). Additional informations to the description of the type
material of Rudolphiella lobosa (type species of the genus) is given. R.
lobosa was not found in L. pati (as reported originally by Riggenbach, 1895)
and has never been found since its original description. The new species
differs from other species of Rudolphiella by its larger scolex, a higher
number of testes and by its distinct egg shape. A diagnosis of Rudolphiella is
given. The following taxonomical actions are introduced: Rudolphiella pira-
catinga (Woodland, 1935) comb. nov. for Monticellia piracatinga Woodland,
1935; Monticellia rugata Rego, 1975 syn. nov. of Rudolphiella piracatinga
(Woodland, 1935). The presence of glandular cells within the scolex’s apex
is observed in all known Rudolphiella species 1.e. R. lobosa, R. myoides, R.
szidati, R. piracatinga, R. piranabu. Rudolphiella, the sole genus within
Rudolphiellinae, is widespread in a monophyletic group of hosts (Calophy-
sus-group sensu de Pinna, 1998) suggesting likely coevolution.
Key-words: Proteocephalidea — Monticelliidae — Rudolphiella szidati sp. n.
— Rudolphiella lobosa — Luciopimelodus pati — coevolution — Argentina.
Manuscript accepted le 9.11.1999
82 ALICIA A. GIL DE PERTIERRA & ALAIN DE CHAMBRIER
INTRODUCTION
Riggenbach (1895) described Corallobothrium lobosum parasite of Luciopi-
melodus pati (Valenciennes, 1840) from Paraguay River, Paraguay. Later, Fuhrmann
(1916) redescribed Riggenbach’s material adding information of proglottides’
transverse sections, and erected a new genus Rudolphiella Fuhrmann, 1916 to allocate
Corallobothrium lobosum Riggenbach, 1895. This species has never been found since
its original description.
Woodland (1934) erected the genus Amphilaphorchis and described two species
Amphilaphorchis piranabu Woodland, 1934 and A. myoides Woodland, 1934, both
parasites of Pinirampus pirinampu (Spix, 1829). A year later Woodland (1935),
concluded that genus Amphilaphorchis Woodland, 1934 is a synonym of Rudolphiella
and erected a new subfamily, the Rudolphiellinae.
The examination of Luciopimelodus pati (Valenciennes, 1840) from Parana river
and de la Plata river, Argentina, the type host of R. lobosa (Riggenbach, 1895) revealed
the presence of a new species of Rudolphiella described herein. Since the original
description of Rudolphiella lobosa (Riggenbach, 1895) did not contain data on some
useful morphological structures, the type material of this species 1s restudied.
MATERIAL AND METHODS
Sixty-two specimens of Luciopimelodus pati (Valenciennes, 1840) were
examined for helminths. Forty intestines were placed in lukewarm water in order to
relax the worms and then fixed in AFA, while others were dissected and fixed directly
in hot 4% formaldehyde solution and were subsequently stored in ethanol 75 % V/V.
Entire tapeworms were stained with Langeron’s alcoholic chlorhydric carmine
(Langeron, 1949), differenciated in acid ethanol, dehydrated in ethanol, cleared in
beechwood creosote or in eugenol, and mounted in Canada balsam. Thick transverse
hand-cutting sections of the proglottides were stained following equivalent procedure.
Two scolices and pieces of strobilas were embedded in paraffin, transversely sectioned
at 12-15 um, stained with Weigert’s hematoxylin and counterstained with 1% eosin B
and mounted in Canada balsam. Eggs were mounted in distillated water for drawing.
Two scolices were prepared for scanning electron microscopy (SEM), they were
dehydrated through a gradual series of ethanol, then put in amyl acetate, critical point
dried and sputtered with gold and photographed with a Zeiss DSM 940 A SEM.
Rudolphiella szidati sp. n. type material was deposited at the Helminthological
Collection of the “Museo Argentino de Ciencias Naturales Bernardino Rivadavıa”,
Buenos Aires, Argentina (MACN), and at the Natural History Museum, Geneva,
Switzerland (MHNG). We also studied syntypes of R. piranabu (Woodland, 1934)
BMNH 1964.12.15.101.107 and syntypes of R. myoides (Woodland, 1934) BMNH
1964.12.15.108.110, (Amaz 74.3), both parasites from Pinirampus pirinampu (Spix,
1829); syntypes of À. piracatinga (Woodland, 1935) BMNH 1964.12.15.206-208,
(Amaz 40) from Calophysus macropterus (Lichtenstein, 1819) from the British
Museum of Natural History, London (BMNH). Amaz = field numbers of Woodland’s
material.
RUDOLPHIELLA SZIDATI SP. N., PARASITE OF LUCIOPIMELODUS PATI FROM ARGENTINA 83
All measurements are given in micrometres, unless otherwise stated, with the
range followed by the mean (m) and the number of measurements (n) in parentheses.
MT = type material. Illustrations were made with the aid of a camera lucida.
RESULTS
Rudolphiella Fuhrmann, 1916
Syn. Amphilaphorchis Woodland, 1934
Diagnosis: Proteocephalidea, Monticelliidae, Rudolphiellinae. Worms of small
or medium size; wrinkle collar-like metascolex; suckers uniloculate with developed
internal circular musculature in their distal margin; apical glandular cells arranged in a
cross situated between the suckers and the apical tegument; internal longitudinal
musculature developed; testes cortical in one layer; vagina posterior or anterior to cirrus
pouch, when anterior ventrally overlapping the cirrus pouch; genital pores irregularly
alternating; ovary medullar with outgrowths in cortex, strongly lobulate; vitelline
follicles cortical, ventral, with a tendency of posterior concentration; uterine primor-
dium and lateral branches medullar; eggs with elongated shell and with embryophores
bearing two elongated polar projections; all species parasites of catfishes of the family
Pimelodidae. Type species: Rudolphiella lobosa (Riggenbach, 1895)
Rudolphiella szidati sp. n. Figs 1, 3-5, 8, 11
Host: Luciopimelodus pati (Valenciennes, 1840) (= Pimelodus pati), common name: patt.
Material studied: Argentina, Provincia de Corrientes, Ciudad de Corrientes, Puerto Italia (Parana
river), holotype MHNG: 26251 INVE, 30.07.1997, paratypes MHNG: 26252 - 26256 INVE,
30.07. to 01.08.1997; Argentina, Provincia de Chaco, Pte General Belgrano (Parana river) (27°
27'S, 58° 50°W), 8 paratypes MACN: 392/1-8; Argentina, Provincia de Buenos Aires, Puerto de
Buenos Aires (de la Plata river) (34° 37°S, 58° 22°W), MACN 392/9-10, 2 paratypes, 06.06.1995;
other material: Argentina, Provincia de Corrientes, Ciudad de Corrientes, Puerto Italia (Parana
river), 24667 - 24674, 27235 INVE, 30-31.07.1997 and MACN 392/11-15.
Site of infection: anterior portion of intestine and rarely in first portion of
medium intestine.
Prevalence: 100%, 62 fishes examined.
Intensity: 2-52; m = 20
Etymology: the new species honours Prof. Lothar Szidat (Buenos Aires).
DESCRIPTION (based on 30 specimens and measurements on 8 specimens):
Monticelliidae, Rudolphielliinae. Testes cortical, situated dorsally and surroun-
ding laterally and ventrally the longitudinal muscular bundles, reaching but not over-
lapping the vitelline follicles (Fig. 4); ovary medullar with projections into the cortical
parenchyma (Fig. 5); uterus medullar (uterine stem and lateral branches). Vitelline
follicles cortical in 2 ventral bands (Figs 1, 4-5).
Medium size worms, flattened dorso-ventrally, total length 16-27 mm (m = 22, n
= 8). Strobila with wrinkles and furrows, acraspedote, comprising about 22-37 (m = 28,
n = 8) proglottides, fast maturation (10-18 immature proglottides, 4-6 mature pro-
glottides and 6-10 gravid proglottides).
84
ALICIA A. GIL DE PERTIERRA & ALAIN DE CHAMBRIER
ROS A) En ae = 2 g &
A
ci 28 68 < EN 7 ; > rte i 2 0. a
= SERI GE Sa DIL > SER = Na = oS
SR OD PRES o Ÿ DU = =
Te POS —
D>
a
RUDOLPHIELLA SZIDATI SP. N., PARASITE OF LUCIOPIMELODUS PATI FROM ARGENTINA 85
Wrinkle collar-like metascolex (in the sense of de Chambrier & Paulino, 1997)
(Fig. 11), encircling the anterior part of the scolex, 1735-2970 in diameter. Metascolex
with apical elongated bottle-shape glandular cells with granular inclusions, arranged in
a cross situated between the suckers and the apical tegument (Fig. 3a); 4 slightly oval
uniloculate suckers. 395-595 (m = 490, n = 19) long and 345-560 (m = 415, n = 19)
wide; presence of internal circular musculature on distal margin of suckers (Fig. 3b):
suckers usually hidden by the collar-like metascolex. The suckers are visible when the
scolex is fixed outside the gut.
Proliferation zone 990-2970 (m = 2005, n = 8) long, 625-1050 (m = 845, n = 8)
wide. Immature proglottides 130-530 (m = 310, n = 26) long, 695-1485 (m = 1080, n =
26) wide; mature proglottides 380-1110 (m = 610, n = 34) long, 925-1780 (m = 1330, n
= 34) wide. Gravid proglottides 495-1500 (m = 965, n = 37) long, 775-1895 (m = 1425,
n = 37) wide. Last proglottis 2145 long, 990 wide.
Internal longitudinal musculature developed, forming a thick bundle of thin
muscular fibres (Figs 4-5). Osmoregulatory canals thick-walled, with anastomoses in
the last third of proglottis. Ventral canals interconnected posteriorly to ovary (Fig. 1).
Ventral canals 20-45 in diameter and dorsal canals 8-30 in diameter. A thin lateral
ventral secondary duct of ventral osmoregulatory canal opens on each posterior side of
the proglottis, forming a vesicle before ending in the tegument. This structure was also
observed in R. lobosa by Riggenbach (1896).
Testes spherical, 50-70 in diameter, 289-365 (m = 325, n = 6), one-layered, in a
compact single field, located in the cortical parenchyma, dorsal, lateral and ventral in
part, not interrupted at cirrus pouch level; overlapping the anterior margin of the ovary
and laterally reaching the posterior end of the proglottis (Fig. 1).
Genital pores alternating irregularly, in 20-46% (m = 30%, n = 34) anterior of
proglottis length. Ejaculatory duct coiled. Cirrus pouch thin-walled, 230-330 (m = 270,
n = 34) long, 60-110 (m = 85, n = 34) wide. Cirrus pouch length occupying 15-32% (m
= 23%, n = 34) of proglottis width. Vas deferens bulky and very coiled, sometimes
reaching body midline, usually reaching anterior margin of mature proglottis,
occupying aproximately one third of mature proglottis length (Fig. 1).
Vagina thick- walled, forming 1-2 loops near the ovary. Vagina posterior (77%)
or anterior (23%) to cirrus pouch; when the vaginal duct is anterior, ventrally
overlapping cirrus pouch, not surrounding the coiled vas deferens as is common. Ovary
medullar, bilobed, strongly lobulate with dorsal and ventral outgrowths, only dorsal
projections reach the cortex. Ovary occupying 43-68% (m = 53%, n = 34) of proglottis
width (Fig. 1).
Fics 1-2
I. Rudolphiella szidati sp. n., holotype 26251 INVE, pregravide proglottis, dorsal view.
2. Rudolphiella lobosa (Riggenbach, 1896), syntype-material, 43/43, pregravide proglottis,
ventral view (coll. Zoological Institute, Neuchatel). The vitelline follicles on the right side are not
entirely represented. Scale-bar, 500 um.
ALICIA A. GIL DE PERTIERRA & ALAIN DE CHAMBRIER
86
iew of scolex, detail of the apical
INVE, apical v
=
D
2467
3a.
idati Sp. n.
Be
b. Rudolphiella s
portion showing the granular cells in a cross shape situated below the tegument. 3b. 27235 INVE,
parasagittal
FIGS 3a-
circular musculature in their distal margin.
oO
section of scolex showing the internal
Scale-bar, 500 um.
RUDOLPHIELLA SZIDATI SP. N., PARASITE OF LUCIOPIMELODUS PATI FROM ARGENTINA 87
ao ae
os
Fics 4-5
Rudolphiella szidati sp. n., paratype, 26253 INVE, transverses sections of a pregravide proglottis.
4. Sections at ovary level; 5. Sections at posterior part of proglottis. Scale-bar, 1000 um.
Vitelline follicles cortical in 2 ventral bands, sometimes reaching anterior and
posterior margin of proglottis, with a tendency to a posterior concentration of follicles,
uninterrupted by cirrus pouch and vagina (Figs 1, 4-5).
Uterine primordium medullar originating from a cylindrical mass of chromophil
cells. Lumen appearing in first mature proglottides. Medullar uterine branches up to
70% (m = 34%, n = 37) of gravid proglottis width, with 9-17 lateral branches opposite
to cirrus pouch and 7-16 on cirrus pouch side.
Eggs with elongated polar projections aproximately of equal size (Fig. 8); thin,
hyaline and spindle-drop shaped external shell. Outer envelope 128-167 (m = 140, n =
12) long, 20-23 (m = 21, n = 12) wide. Inner envelope consisting in bilayered embryo-
phore, with nucleate envelope 16-21 (m = 20, n = 12) long, 10-16 (m = 14, n= 12) wide
and external layer 40-50 (m = 45, n = 12) long, 18-21 (m = 20, n = 12). Oncosphere
slightly oval, 10-13 (m = 11, n = 12) long, 7-13 (m = 10, n = 12) wide; oncosphere
hooks, 3-6 (m = 4, n = 17); According to Swiderski (1994), we interprete the described
structures as: 1. Shell; 2. Outer envelope; 3. Inner envelope consisting in bilayered
embryophore with external layer much bigger than nucleate envelope; 4. Oncospherical
membrane rarely visible; 5. Oncosphere.
88 ALICIA A. GIL DE PERTIERRA & ALAIN DE CHAMBRIER
Rudolphiella lobosa (Riggenbach, 1895) Fuhrmann, 1916 Fig. 2
syn. Corallobothrium lobosum Riggenbach, 1895; Ephedrocephalus lobosum (Riggenbach,1895)
Mola, 1906.
Host (according to Riggenbach, 1895): Pimelodus pati (= Luciopimelodus pati (Valenciennes,
1840), common name: pati.
Material examined: Paraguay, Paraguay river, type-material, slides 43/43-44 (Collection of the
Institute of Zoology, Neuchatel, deposited in the MHNG), collected by Ternetz, 01-02.1894.
Site of infection: intestine.
DESCRIPTION:
Strobila acraspedote, with wrinkles and furrows; worms of medium size 17-22
mm long, with fast maturation. Scolex with apical glandular cells arranged in a cross,
situated between the suckers and the apical tegument, presence of a well developed
metascolex, 1025-1350 wide. Suckers uniloculate 205-245 in diameter, with free distal
muscular sphincter. Proliferation zone about 800 long. Longitudinal muscular fibers
developed all around the proglottis.
Testes in one layer in transverse sections, in a single field, sphaerical 50-65 in
diameter, 194-219 (m = 206; n = 3) (53 in original figure of Riggenbach (1896), 150-
200 according to Riggenbach (1896), 220 according to Fuhrmann (1916)) in number.
Testes in cortical parenchyma, dorsal, lateral and ventral in part, not interrupted at
cirrus pouch level; overlapping anterior margin of ovary and laterally reaching posterior
end of the proglottis (Fig. 2).
Genital pores alternating irregularly, in 21-32% (n = 6) of proglottis length.
Cirrus pouch thin-walled, 170-205 long. Cirrus pouch length occupying 16-22% (n= 4)
of proglottis width. Vas deferens not reaching body midline in mature proglottis.
Vagina posterior (53%) or anterior (47%) to cirrus pouch, with inconspicuous
terminal muscular sphincter. Ovary medullar, bilobed, strongly lobed and bearing
dorsal projections penetrating the cortex; occupying 60-72% of proglottis width.
Vitelline follicles distributed cortically and ventrally, in two lateral bands, sometimes
reaching anterior and posterior margin of proglottis, with a tendency of posterior
concentration, uninterrupted by cirrus pouch and vagina.
Uterine primordium like a medullar cylindrical mass of chromophil cells.
Lumen appearing from the first mature proglottides until formation of eggs. Uterine
branches occupying up to 56% of proglottis width, with 8-12 lateral branches on each
side, lor 2 layered. Eggs with two polar projections.
Remarks: The new species belongs to Rudolphiella Fuhrmann, 1916 based on
the cortical distribution of the testes and vitelline follicles, on the medullar position of
uterus, as well as on the medullar and partially cortical location of the ovary (Rego,
1994). Four species of Rudolphiella are present: R. lobosa (Riggenbach, 1895) recorded
in Luciopimelodus pati: R. myoides (Woodland, 1934) and R. piranabu (Woodland,
1934), both parasites of Pinirampus pirinampu; Rudolphiella piracatinga (Woodland,
1935), parasite of Calophysus macropterus. Rudolphiella cf. lobosa de Chambrier &
Vaucher, 1999 parasite of Megalonema platanum is also considered.
RUDOLPHIELLA SZIDATI SP. N., PARASITE OF LUCIOPIMELODUS PATI FROM ARGENTINA 89
Rudolphiella szidati sp. n. differs from all described species in the genus by the
size of metascolex and by the testes number (Table 1). Furthermore, in R. szidati, (with
R. lobosa and R. cf. lobosa) testes laterally reaching the posterior end of proglottis,
while in R. myoides and R. piranabu the number of testes decreases and they do not
Fics 6-10
Rudolphiella spp., Eggs drawn in distilled water. 6. Rudolphiella piranabu (Woodland, 1934),
25129 INVE; 7. Rudolphiella myoides (Woodland, 1934), 24712 INVE; 8. Rudolphiella szidati
sp. n., 24670 INVE; 9. Rudolphiella piracatinga (Woodland, 1935), 19650 INVE; 10.
Rudolphiella cf. lobosa, 22352 INVE (de Chambrier & Vaucher, 1999, fig. 95). Several eggs are
not totally ripe. Abbreviations: eb = bilayered embryophore, oe = outer envelope, on =
oncosphere, s = shell. Scale-bar, 50 um
90 ALICIA A. GIL DE PERTIERRA & ALAIN DE CHAMBRIER
reach the posterior end of proglottis (de Chambrier & Vaucher, 1999; Pavanelli &
Machado dos Santos, 1992; Riggenbach, 1895; Woodland, 1934, 1935).
Within the genus Rudolphiella, the eggs have a similar pattern with elongated
poles, R. szidati (Fig. 8), R. piracatinga (Fig. 9), R. cf. lobosa (Fig. 10) and R. piranabu
(Fig. 6) possess poles of similar size, but the shape is different in the 4 taxa; in R.
myoides, one pole is shorter than the other (Fig. 7). The oncospheres of the 4 known
species are similar in size (9-13), the outer envelope is difficult to compare among
species based on uncertainties in the descriptions by different authors.
DISCUSSION
As we observed some more characters which are shared by all Rudolphiella
species (i.e. presence of glandular cells within the apex, vagina anterior/posterior, shape
of eggs), we gave above a new diagnosis of the genus.
Luciopimelodus pati is a common dweller fish of the Paraguay river, Bermejo
river, Parana river, Uruguay river, Carcarana river and de la Plata river. The distribution
of 3 Pimelodidae sharing the same common name (« pati »), Megalonema platanum
(Günther, 1880), Pinirampus pirinampu (Spix, 1829) and Luciopimelodus pati
(Valenciennes, 1840) is overlapping (Ringuelet er al., 1967). Since we were unable to
find the species of parasites described by Riggenbach (1895) (neither R. lobosa nor
Proteocephalus fossatus), from L. pati, we suspect that the host studied by Riggenbach
was not Luciopimelodus pati, but possibly Megalonema platanum; this host was studied
for cestodes by de Chambrier & Vaucher (1999) and it was parasited by Rudolphiella
cf. lobosa.
Rego (1975) described Monticellia rugata from Calophysus macropterus in
Amazonia. This cestode is a mixture of the two species Nomimoscolex piracatinga
Woodland, 1935 and Monticellia piracatinga Woodland, 1935. The figures 44, 45, 47,
49 and 50 belong to Monticellia piracatinga and figures 46 and 48 belong to Nomi-
moscolex piracatinga. This latter species was transfered to the genus Monticellia under
the name Monticellia amazonica by de Chambrier & Vaucher (1997). «Piracatinga» is
the vernacular name given to Calophysus macropterus in Amazonia and to
Luciopimelodus pati in high Parana river in Brazil. Fowler (1951) and Ringuelet er al.
(1967) placed L. pati into the Parana bassin, but not in the Amazon bassin. We think
that Woodland could be mistaken in attributing the name piracatinga to L. pati in
Amazonia. Our recently collected material in Amazonia from Calophysus macropterus
fits with the type material of the two species described by Woodland (M. piracatinga
and N. piracatinga) and seems to confirm our hypothesis. Monticellia rugata Rego,
1975 is thus a junior synonym of Monticellia piracatinga. However, the study of type
material of M. piracatinga Woodland, 1935 revealed that this species belongs to the
genus Rudolphiella, as it is confirmed by the recent collected material from Calophysus
macropterus in the Amazon and becomes Rudolphiella piracatinga (Woodland, 1935)
comb. nov.
So far, numerous proteocephalideans possess unicelullar glands in the scolex (de
Chambrier ef al., 1992, 1996; de Chambrier & Vaucher, 1997, 1999; Gil de Pertierra,
RUDOLPHIELLA SZIDATI SP. N., PARASITE OF LUCIOPIMELODUS PATI FROM ARGENTINA 9]
Fic. 11
Scolex, Rudolphiella szidati sp. n., 24669 INVE. Scale-bar, 200 um.
1995; Gil de Pertierra & Viozzi, 1999; Scholz et al., 1998, 1999; Stoitsova et al., 1995).
These glands are also present in all known species of Rudolphiella in high number and
arranged in a cross, with granular inclusions. Zd“arska & Nebesarova (1999) give a
detailed description of the unicellular glands situated under the apex of the scolex in
Proteocephalus macropterus.
The presence of R. myoides and R. piranabu was confirmed in the host Pini-
rampus pirinampu from Amazon river by one of the authors (A. de Chambrier), and R.
piranabu was confirmed by Pavanelli & Machado Dos Santos (1992) in the same host
from Parana river, Brazil.
Brooks (1984, 1995) already demonstrated, in phylogenetic analyses based on
morphological characters, the monophyly of three Rudolphiella species (R. lobosa, R.
myoides and R piranabu) based on the presence of elongated egg. However, his 1995
analysis showed some homoplasious characters and is consequently somewhat less
ALICIA A. GIL DE PERTIERRA & ALAIN DE CHAMBRIER
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RUDOLPHIELLA SZIDATI SP. N., PARASITE OF LUCIOPIMELODUS PATI FROM ARGENTINA 93
supported. We observed that all Rudolphiella species shown some more synapo-
morphies: presence of glandular cells with granular inclusions arranged in a cross
situated between the suckers and the tegument, eggs with elongated shell, embryo-
phores bearing two polar projections, vitellines follicles cortical in two ventral bands
situated sub-laterally, vagina anterior/posterior. Those synapomorphies shared by all
Rudolphiella species emphasize the homogeneity of this group and enforced the
Brooks’ opinion on the monophyly of this genus.
Stewart (1986) stated that Luciopimelodus pati, Pinirampus pirinampu and
Calophysus macropterus belong to a monophyletic group called the «Calophysus
group», within the Pimelodidae, opinion confirmed by Lundberg ef al. (1991) and more
recently by de Pinna (1998). On the other hand, Megalonema shares some charac-
teristics with the Calophysus group, and other groups closely related to Diplomystidae
(primitive catfishes). In the light of our studies, the presence of Rudolphiella ct. lobosa
(de Chambrier & Vaucher, 1999) from Megalonema platanum confirms the relationship
of M. platanum with the Calophysus group.
In conclusion, the sole genus Rudolphiella is widespread among the genera of
the «Calophysus group» we studied, including Megalonema platanum. It is, worthwhile
to emphasize that it could be a likely case of coevolution between related species of
hosts and their parasites.
ACKNOWLEDGEMENTS
We are grateful to Dr. C. Vaucher for helpful discussions and for inviting the
first author to the Natural History Museum, Geneva; to the rewievers for constructive
critisism; to the Municipal Departement of Cultural Affaires, Geneva, Switzerland, for
supporting her stay at the Department of Invertebrates of the Museum in 1998. Her visit
was also supported by the Secretaria de Recursos Naturales y Desarrollo Sustentable de
la Nacion, Buenos Aires, Argentina. We thank the staff of the Centro de Ecologia
Aplicada del Litoral, Corrientes, Argentina for assisting our stay in the field during
1997 and particularly to Mr. L. Benetti, Prof. J. J. Neiff, Dr. M. Hamann and Dr. A.
Kehr. We are thankful to Dr. E. A. Harris, at the British Museum of Natural History, for
the loan of specimens.
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È
| | |
RA ABR Talarico
REVUE SUISSE DE ZOOLOGIE, 107 (1): 97-106; mars 2000
Two new species of the genus Suffasia from Sri Lanka
(Araneae: Zodariidae)
Suresh P. BENJAMIN! & Rudy JOCQUE?
l'Institut für Natur-, Landschafts- und Umweltschutz der Universität Basel (NLU),
Abteilung Biologie, St. Johanns-Vorstadt 10
CH-4056 Basel, Switzerland.
2 Koninklijk Museum voor Midden-Afrika, B-3080 Tervuren, Belgium.
Two new species of the genus Suffasia from Sri Lanka (Araneae:
Zodariidae). - Two new species of the genus Suffasia Jocqué, 1991 are
described. S. mahasumana sp. n. 1s known from both sexes and is related to
S. tumegaster Jocqué, 1992. S. attidiva sp. n., known only from two female
specimens, may be related to S. tigrina (Simon, 1893). S. attidiya Sp. n. is
found in diverse habitats, S. mahasumana sp. n. is confined to cloud forests
in the central highlands of Sri Lanka. This is the first record of Suffasia
from Sri Lanka, other species in this genus are known only from southern
India and Nepal.
Key-words: Araneae - Zodariidae - Suffasia - tropical montane cloud forests
- Sri Lanka.
INTRODUCTION
The genus Suffasia was established for two species from southern India. The
type species, S. tigrina (Simon, 1893) from Kodaikanal, Tamil Nadu, and an undescri-
bed species from the same locality (Jocqué, 1991). S. tumegaster Jocqué, 1992, from
Kathmandu, Nepal, was added later. Considering the affinities of the faunas of India
and Sri Lanka it was to be expected that members of Suffasia or at least of closely
related genera might be discovered on the island.
During recent field work in Sri Lanka two undescribed species were collected.
The first one is from the Knuckles Range, consisting of a number of remnant patches of
primary tropical montane cloud forests, a type of vegetation that was once common in
the central highlands of Sri Lanka. Most of these forests were cleared for tea plantations
during the British colonial period. The second species was collected in fragmented
marshland situated on the outskirts of Colombo. On both localities the specimens were
obtained by beating shrubs and small trees which is an unusual way to collect zodariids
as they are considered soil dwelling spiders except for the representatives of the
Storenomorphinae (Jocqué, 1991). In the Bellanwila-Attidiya sanctuary a marshy area
near Colombo, shrubs are scattered around shallow water ponds, marshes and sea-
Manuscript accepted 21.10.1999
98 SURESH P. BENJAMIN & RUDY JOCQUE
sonally flooded grassland. A female of the second new species was collected by the
same method in Kalugala, Labugama Forest Reserve, a fragmented remnant of tropical
lowland rain forest, some 40 km away from the former locality.
METHODS
Structures were examined in temporary mounts embedded in glycerine. Vulvae
were Cleared with trypsin (0.1% trypsin, 0.1% CaCl,, in 0.05M tris-buffer, pH 7.6). All
drawings were made with a Nikon Labophot-2 and a Nikon SMZ-U microscopes with
drawing tube. Measurements are in mm. Structures examined with the scanning elec-
tron microscope (PHILIPS XL30 FEG ESEM) were critical point dried, stud-mounted
and sputter coated for observation and photography. Specimens examined are deposited
in the “Muséum d’histoire naturelle, Geneve” (MHNG) and the “Naturhistorisches
Museum, Basel” (NMB).
Abbreviations used in the text and figures: AER anterior eye row; ALE anterior
lateral eyes; AME anterior median eyes; CD copulatory duct; CF cymbial flange; CO
copulatory opening; E embolus; FD fertilisation duct; PER posterior eye row; PLE
posterior lateral eyes; PME posterior median eyes; TA tegular aphophysis.
Suffasia mahasumana sp. n. Figs 1-7, 13-21
Holotype 3: Sri Lanka, Central Province, Knuckles Range, Deenston (approximately 7° 19° N,
80° 51° E), 1100 m, 11 March 1998. Leg. Suresh P. Benjamin (MHNG).
Paratypes: 1%, 11 March 1998 (MHNG): 14,19 12 March 1998 (NMB); further as holotype.
Etymology: Named after god Maha-Sumana, protector of the hill country in Sri Lanka.
Noun in apposition.
Diagnosis: S. mahasumana is closely related to S. tumegaster; the male can be reco-
gnised by the absence of a dorsal spike on the palpal tibia, the shape of the dorsal
cymbial flange which is flat and evenly rounded in the latter, swollen, curved upwards
and concave in the former; the female differs from that of S. tumegaster by the presence
of a roughly rectangular plate in the anterior part of the epigyne.
Description: Male (holotype). Colouration and markings: carapace dark yellow-brown,
with dark reticulations on anterior part and with U-shaped dark marking in front of
fovea (Fig. 14). Chelicerae and sternum dark yellow, lighter than carapace. Dorsum of
opisthosoma uniform darkgrey, venter white, without markings. Legs light yellow with
dark dorsal markings. AER almost straight, PER slightly procurved, all eyes circular,
AME 1.5 times their diameter apart from each other and at about the same distance
from ALE. PME 2 times their diameter apart and about the same distance from PLE.
ALE = PLE = PME> AME. Clypeus height 6 times the diameter of ALE. Chilum
present. Chelicerae not fused, with double-tooth on promargin. Labium triangular.
Sternum sub triangular, with spike-like extensions projecting towards base of coxae.
Leg formula 4132, 2 spines dorsally on femora I-IV. Tibiae with flattened incised hairs
(Fig. 16, FIS); and ventral tuft of metatarsal preening brush with chisel-shaped hairs
(Figs 16, 17, CH; see also Jocqué, 1991: figs 6, 8, 11, 12). Femoral organ (Fig. 20)
present on each leg. Trichobothrium base with concentric ridges (Fig. 15).
TWO NEW SUFFASIA FROM SRI LANKA 99
Palp (Figs 1-4): Tibia with stout, sharp, strongly tapered retrolateral apophysis,
pointing outwards. Cymbium strongly narrowed in dorsal view, with swollen, upwards
curved, dorsolateral flange (Fig. 1, CF), extending lateral cymbial concavity, carrying
some sensorial hairs in superior part; embolus fairly short, stout, originating on
Measurements: total length: 2.3; carapace length: 1.3; carapace width: 1.0. Legs:
I Il III IV
femur 13] 0.9 1.0 iol
patella 0.2 0.2 0.3 0.3
tibia 1.0 0.7 0.7 1.0
metatarsus 1] 0.9 1.0 LS
tarsus 0.5 0.4 0.4 0.5
total 3.9 3.1 3.4 4.4
posterior part of tegulum separated from main part by shallow groove; tegular
apophysis, short, stout, sharp, pointing out and forwards (Figs 1, 2, TA).
Female. Colouration and markings: Similar to male but lighter. Different by
possessing dorsal opisthosomal markings as in Fig. 13; venter white. Palp with conical
tarsus, longer than tibia (Fig. 19). Morphology further as in male.
Epigynum and vulva (Figs 5-7): Simple brown plate in anterior part; internal structure
visible through thin tegument; copulatory openings in front hidden by plate; short
copulatory ducts lead to thick-walled spermathecae, triangular in dorsal view.
Measurements: Total length: 3.0; carapace length: 1.3; carapace width: 1.0. Legs:
I II III IV
femur 0.8 0.8 0.9 PSI
patella 0.3 0.3 0.4 0.4
tibia 0.8 0.7 0.6 0.8
metatarsus 0.9 0.7 0.8 1.2
tarsus 0.5 0.4 0.4 0.5
total 33 2.9 Bell 4.0
Affinities: Suffasia is defined by the presence of a chilum and promarginal cheliceral
teeth, dark reticulation (“network pattern” sensu Jocqué, 1992), female palp with a long
conical tarsus, femoral organ with simple setae on all legs, legs with flattened incised
hairs and metatarsal preening brush consisting of chisel-shaped hairs (Jocqué, 1991,
1992). The present species clearly agrees with these characters and can thus be
attributed to Suffasia. Yet the shape of the male palpal cymbium casts some doubt on
this attribution. Although one of the main characteristics of Asceua Thorell, 1887 is
exactly the strongly narrowed cymbium, there are a number of characters that exclude
the incorporation of the present species in it: representatives of that genus do indeed
SURESH P. BENJAMIN & RUDY JOCQUE
100
Fics 1-4. Suffasia mahasumana sp. n. 1. Male palp, retrolateral view. 2. Ditto, ventral view. 3.
Ditto, dorsal view. 4. Ditto, prolateral view. CF cymbial flange; E embolus; TA tegular
apophysis. Scale line: 0.2 mm.
TWO NEW SUFFASIA FROM SRI LANKA 101
7
Fics 5-7. Suffasia mahasumana sp. n. 5. Female epigynum, ventral view. 6. Vulva, ventral view.
7. Ditto, dorsal view. CD copulatory duct; FD fertilisation duct. Scale line: 0.1 mm.
102 SURESH P. BENJAMIN & RUDY JOCQUE
lack teeth, have a uniform dark carapace and lack femoral organs. Yet, both S. maha-
sumana and S. tumegaster possess an epigynum that is quite different from what has
been described for Suffasia. It might prove necessary to erect a new genus for these
species if the male of type species of Suffasia appears equally different which was
already recognised by Jocqué (1992).
Distribution: Known only from the type locality.
Suffasia attidiya sp. n. Figs 8-12
Holotype 2: Sri Lanka, Western Province, Colombo, Bellanwila-Attidiya (approximately
6°50°N, 79°54’E), mean elevation 0.6 m asl, 22 February 1998, (MHNG).
Paratype 2: Sri Lanka, Western Province, Kalugala, Labugama Forest Reserve, 3 August 1996,
ca. 10 m (NMB). All specimens leg. Suresh P. Benjamin.
Etymology: Named after the type locality. Noun in apposition.
Diagnosis: The epigyne of S. attidiva differs from that of S. tigrina by the course of the
copulatory ducts which run directly inwards in the latter, outwards thence inwards in
the new species. S. mahasumana is clearly different by the presence of a plate in the
epigyne.
Description: Female (holotype). Colouration and markings: prosoma dorsally dark
yellow-brown, with dark markings in front of fovea (Fig. 12). Chelicerae and sternum
dark yellow lighter than dorsal prosoma. Dorsum of opisthosoma with markings as in
Fig. 11, venter white. Legs light yellow with dark dorsal markings. AER slightly pro-
curved, PER procurved, all eyes circular, AME 1.5 times their diameter apart from each
other, 0.5 times from ALE. PME 2 times their diameter apart and 2.5 times from PLE.
ALE = PLE = PME > AME. Clypeus height 6 times the diameter of ALE. Chilum
present; chelicerae not fused, with double tooth on promargin. Labium triangular. Ster-
num sub triangular, with spike-like extensions projecting towards base of coxae. Palp
with conical tarsus, longer than tibia. Flattened incised hairs on tibiae; ventral tuft of
metatarsal preening brush with chisel-shaped hairs.
Epigynum and vulva (Figs 8-10): Epigynum simple, anterior sclerotized border with
CO situated laterally. Thick-walled copulatory ducts straight and close to each other in
posterior part, leading to small globular rectptacula, with thick walls. FD as in Fig. 9.
Male. Unknown.
Affinities: As zodariid genera are mainly diagnosed on male palpal morphology
(Jocqué, 1991, 1992) the placement of this new species of Suffasia might appear ambi-
guous. However, the similarities of the epigynum of the present species and of the type
species are so striking that there is little doubt that they are congeneric. In both cases
the internal structure is simple with lateral CO, strongly sclerotized CD with a partly
parallel course and roughly oval spermathecae.
Fics 8-14. Suffasia attidiva sp. n. (8-12). Suffasia mahasumana sp. n. (13, 14). 8. Female
epigynum, ventral view. 9. Vulva, ventral view. 10. Ditto, dorsal view. 11. Female opisthosoma,
dorsal view. 12. Female prosoma and right palp, dorsal view. 13. Female, dorsal view. 14. Male,
dorsal view. CO copulatory opening; FD fertilisation duct. Scale lines: 0.1 mm (8-10), 2.0 mm
(11-14).
103
ASIA FROM SRI LANKA
£
TWO NEW SUFF
104 SURESH P. BENJAMIN & RUDY JOCQUE
Fics 15-18. Suffasia mahasumana sp. n., SEM micrographs. 15. Base of trichobothrium. 16.
Preening brush on distal metatarsus of leg I, lateral view. 17. Ditto, detail. 18. Tip of leg I, lateral
view. CH chisel-shaped hairs, FIS flattened incised hairs. Scale lines: 0.001 mm (1), 0.01 mm
(16-18).
TWO NEW SUFFASIA FROM SRI LANKA 105
Fics 19-21. Suffasia mahasumana sp. n., SEM micrographs. 19. Chelicerae and palps of female,
frontal view. 20. Femoral organ, on leg I. 21. Chemosensitive hair on metatarsus. Scale lines:
0.003 mm (20), 005 mm (21), 0.1 mm (19).
106 SURESH P. BENJAMIN & RUDY JOCQUE
Measurements: Total length: 2.6; carapace length: 1.3; carapace width: 1.0. Legs:
I II III IV
femur 0.6 0.5 0.6 0.7
patella 0.1 0.1 0.2 0.2
tibia 0.5 0.4 0.4 0.6
metatarsus 0.6 0.5 0.5 0.8
tarsus 0.4 0.3 0.3 0.4
total DID, 1.8 2.0 DOT
Distribution: Known from Bellanwila-Attidiya sanctuary and Kalugala, Labugama
Forest Reserve.
DISCUSSION
The placement of the two new species in the genus Suffasia is in accordance
with the current definition of the genus. However the relationships proposed here
should be re-analysed when additional material, most importantly the male of S. tigrina
is discovered.
In his revision of the Zodariidae Jocqué (1991) considered the subfamilies
Zodariinae and Storeninae to be monophyletic. This hypothesis was based on presumed
autapomorphies such as the presence of a femoral organ and flattened incised hairs for
the Zodariinae and chisel-shaped hairs on the metatarsal preening brush for Storeninae.
The discovery of S. tumegaster, which possesses a combination of all these characters,
led to an amalgamation of these two subfamilies (Jocqué, 1992). S. mahasumana sp. n.
which also possesses these three characters, further confirms his combination of both
subfamilies.
The discovery of the new taxa extends the previously known distribution (Nepal,
India) of the genus Suffasia southwards to Sri Lanka.
ACKNOWLEDGEMENTS
We thank Dr. Peter Schwendinger (MHNG) for critical review of the manuscript
and for his encouragement during this study. Part of this work was done in the course of
the masters thesis of the first author at the University of Innsbruck. He is grateful to Dr.
K. Thaler for providing research facilities there. We also thank Marcel Diiggelin
(Raster lab of the Unviersity of Basel) for help with SEM work, Mr. D. Benjamin
(Colombo) for accompanying the first author on collecting trips to the study area and
Mr. A. H. Sumanasena (Department of Wild Life Conservation, Colombo) for pro-
viding a research permit.
REFERENCES
JOCQUÉ. R. 1991. A generic revision of the spider family Zodariidae (Araneae). Bulletin of the
American Muesum of Natural History 201: 1-165.
JOCQUÉ, R. 1992. A new species and the first males of Suffasia with a redelimitation of the
subfamilies of the Zodariidae (Araneae). Revue suisse de Zoologie 99: 3-9.
REVUE SUISSE DE ZOOLOGIE 107 (1): 107-122; mars 2000
Diversite du zoobenthos dans 47 rivieres du canton de Vaud:
tendance 1989 - 1997
Claude LANG
Conservation de la faune, Marquisat 1, CH-1025 St-Sulpice, Suisse.
claude.lang@sffn.vd.ch
Diversity of zoobenthos in 47 rivers of western Switzerland: the 1989 -
1997 trend. - Three surveys of benthic invertebrates were made between
1989 and 1997 in 165 sites located in 47 rivers of western Switzerland
(canton of Vaud). Eight descriptors derived from the total number of taxa
and the number of taxa intolerant of pollution (Plecoptera, Heptageniidae,
and Trichoptera with a case) were used to describe the patterns of benthic
diversity. In the three surveys, the diversity increased with the altitude of
sampling sites. This trend reflected the upward decrease of human impact
on the rivers and their watersheds. The diversity increased between 1989
and 1997 because of better meteorological conditions (i.e. more water in
the rivers) coupled with the increased efficiency of sewage treatment
plants. But the diversity remained low in several small rivers, probably
because of chronic pollutions by pesticides.
Key-words: diversity - indicator species - river - water quality - zoo-
benthos.
INTRODUCTION
Depuis la signature de la convention sur la diversité biologique a Rio de Janeiro
en 1992, la conservation et la restauration de la diversité des especes dans l’espace et le
temps constituent un des buts fondamentaux de l'écologie appliquée (Wright er al.,
1998). Dans le même ordre d'idée, la diversité des communautés d’invertebres qui colo-
nisent le fond des cours d'eau (zoobenthos) est utilisée depuis le début du siècle pour
caractériser l’état du milieu: elle diminue en fonction de l'intensité des perturbations
subies, que celles-ci soient d'origine humaine ou naturelle (Hellawell, 1986).
Cette diminution de la diversité est particulièrement marquée chez les plé-
copteres, certains éphéméroptères et les trichopteres à fourreau d’où l’utilisation de ces
groupes comme indicateurs (Fore ef al., 1996). Dans les rivieres du canton de Vaud par
exemple, la diversité des plécoptères a fortement diminué entre 1945 et 1982 à basse
altitude alors qu'elle ne changeait guère dans les rivières de montagne, restées proches
de l'état naturel (Aubert, 1984). Ce gradient de diversité amont-aval reflète l’augmen-
tation des impacts d'origine humaine à basse altitude (Lang & Reymond, 1993, 1995).
Manuscrit accepté le 28.07.1999
108 CLAUDE LANG
Entre 1982 et 1997, l'épuration des eaux s'est généralisée et les performances des
stations d’épuration se sont améliorées dans le canton de Vaud (Fiaux er al., 1998). De
ce fait, les quantités de matiere organique et de nutriments déversées dans les cours
d'eau ont diminué. La diversité du zoobenthos devrait donc augmenter en réponse a
cette amélioration de la qualité chimique de l’eau.
Il ne faut toutefois pas oublier que le bassin versant et la rivière qui le draîne,
forment un tout: ce qui affecte l'un, modifie l’autre (Hynes, 1975). Du fait de cette
relation étroite, le caractere de moins en moins naturel des bassins versants, surtout a
basse altitude, pourrait empécher la restauration de la diversité, méme si la composition
chimique de l'eau redevient normale, tout au moins en termes d’éléments majeurs
(matiere organique, phosphore et azote); les micropolluants constituant en effet un
problème a part (Corvi & Kim-Heang, 1997; Vioget & Strawczynski, 1997).
Pour interpréter l'évolution a long terme du zoobenthos, il faut également tenir
compte de la météorologie (Allan, 1995). Par exemple, une sécheresse de longue durée
va réduire le débit des cours d'eau et augmenter ainsi la concentration des polluants,
donc leur impact sur le zoobenthos. Faute d'une dilution suffisante, les capacités d'auto-
épuration naturelles de certains cours d'eau peuvent étre completement dépassées si
l’épuration des eaux n’a pas été conçue en fonction de conditions extrêmes. Ces
dernieres pourraient s’ observer plus fréquemment si le climat venait a changer rapide-
ment sous l'influence de l’effet de serre. Signalons également que des crues très vio-
lentes, comme une longue sécheresse, peuvent durablement diminuer la diversité du
zoobenthos en bouleversant le lit des cours d’eau (Allan, 1995).
La présente étude analyse l’évolution de la diversité du zoobenthos dans les
rivieres vaudoises sur la base de trois campagnes de prélevements effectuées entre 1989
et 1997. Son but consiste a mettre en évidence l’effet sur le zoobenthos des mesures
d'assainissement prises, ceci en fonction de l’altitude et des conditions météorologiques
observées au cours de cette période.
STATIONS ET METHODES
Le zoobenthos des rivieres vaudoises est étudié depuis 1982 (Lang & Reymond,
1995). Cependant le réseau de surveillance, sous sa forme actuelle, n'a été mis en place
qu'à partir de 1989. De ce fait l'analyse présentée ici se concentre sur une période ou les
données sont parfaitement comparables entre elles: 165 stations de prélevements
étudiées au cours de trois campagnes effectuées entre 1989 et 1997 dans 47 rivières.
Les 47 rivières étudiées (Fig. 1) sont réparties en trois groupes d'après la région
où chacune d'elles prend sa source: (1) le Jura et l'ouest du canton de Vaud, (2) le Jorat
et le centre du canton, (3) les Préalpes et les Alpes. La Broye, I’ Arbogne et la Mionne
sont rattachées aux rivieres du Jorat en raison de leur ressemblance avec celles-ci. Dans
le Jura, 20 rivières et 75 stations de prélèvement ont été visitées en 1990, 1993 et 1996
(Lang, 1997); dans le Jorat, 15 riviéres et 47 stations visitées en 1991, 1994 et 1997
(Lang, 1998); dans les Alpes, 12 riviéres et 43 stations visitées en 1989, 1992 et 1995
(Lang, 1996). Seules les rivières et les stations visitées au cours des trois campagnes
sont incluses dans la présente étude.
DIVERSITE DU ZOOBENTHOS DANS 47 RIVIERES 109
Fic. |
Localisation et numéros d'identification des 47 rivieres vaudoises visitées entre 1989 et 1997.
1 Doye 11 Saubrette 21
2 B. de Nyon 12 B. Morges 22
3 Asse 13 Morges 23
4 Promenthouse 14 Venoge 24
5 Colline 15 Veyron 25
6 Cordex 16 Nozon 26
7 Serine 17 Orbe Dil
8 Dullive 18 Mujon 28
9 Aubonne 19 Arnon 29
10 Toleure 20 Baumine 30
Talent
Buron
Mentue
Sauteru
Petite Gläne
Broye
Arbogne
Lembe
Cerjaule
Mérine
31 Bressonne 41 Gryonne
32 Carrouge 42 P. Gryonne
33 Grenet 43 Avançon
34 Mionne 44 Hongrin
35 Forestay 45 Sarine
36 Veveyse 46 Flendruz
37 B. Clarens 47 Torneresse
38 B. Montreux 1 - 20 Jura
39 Tinière 21 - 35 Jorat
40 Grande Eau 36 - 47 Alpes
Les 165 stations de prélèvements étudiées ont été choisies parce qu’elles pré-
sentent des caractéristiques communes qui les rendent comparables entre elles: pour la
plupart, elles sont localisées dans la zone à truite de Huet et le rithron d’Illies, donc
110 CLAUDE LANG
TABLEAU 1. Calcul de la valeur de l'indice RIVAUD à partir du nombre total de taxons (NT) et du
nombre de taxons sensibles aux pollutions (NTS). La valeur de RIVAUD se lit à l'intersection de
la ligne renfermant la valeur de NT et de la colonne ou se trouve la valeur de NTS. Exemple: NT
= 19, NTS = 7, RIVAUD = 11. Source: Lang & Reymond (1995).
NTS
NT 0 I 2 3 4 5 6-7 8-9 10-11 12-13 14-25
1-4 I 2 3 4 5 - - - - - -
5-8 2 3 4 5 6 7 8 9 - - -
9-12 3 4 5 6 7 8 9 10 ll 12 -
13-16 4 5 6 7 8 9 10 ll 2 13 14
17-20 5 6 7 8 9 10 1] 12 13 14 15
21-24 6 7 8 9 10 1] 12 3 14 15 16
25-28 Ti 8 9 10 1] 12 13 14 [es 16 IM
29-32 8 9 10 1] 12 13 14 15 16 17 18
33-36 9 10 Il 12 13 14 15 16 17 18 19
37-50 10 1] 12 13 14 15 16 17 18 19 20
Qualité biologique mauvaise (RIVAUD | - 5), faible (6 - 9), médiocre (10 - 11), moyenne (12 -
14), bonne (15 - 20).
dans une écorégion relativement homogene (Ribaut, 1966); toutes renferment des
surfaces de cailloux balayées par le courant (zone de rapides) d’au moins 20 m? ce qui
permet de prélever toujours dans le méme substrat; aucune n'est directement exposée
aux rejets polluants d’une station d'épuration ou d'un important égout: elles représentent
donc la situation générale de la rivière en l’absence de pollutions ponctuelles bien
marquées.
Dans le Jura et le Jorat, chaque station est visitée à deux reprises chaque année
d'étude: la première fois en janvier-février, la seconde fois en mars-avril, c'est-à-dire
pendant l’étiage d’hiver et avant la crue de printemps. Dans les Alpes, la premiere
campagne s'effectue en février-mars pendant l'étiage hivernal, la deuxième en juin
après la crue de printemps et la troisième en septembre en période de basses eaux. Trois
visites par année sont nécessaires dans les Alpes. Certaines stations ne sont en effet pas
accessibles en hiver à cause de la neige et la glace, en juin à cause des hautes eaux et en
septembre parce que le niveau d’eau est trop bas pour prélever. De plus, du fait du
caractère torrentiel de ces cours d'eau, l’abondance de la faune y est souvent très faible
ce qui fait qu’une visite peut ne pas être représentative.
Au cours de chaque visite, six coups de filet sont donnés dans six différentes
zones de cailloux (rapides) de la station, correspondant chacune à une surface prélevée
d'environ 0.1 m2. Le filet est posé sur le fond, son ouverture (20 cm x 20 cm) face au
courant, de manière à ce que celui-ci y entraîne les invertébrés délogés en remuant le
substrat avec le pied. Tous les invertébrés récoltés dans ces six coups de filet consti-
tuent un prélèvement qui est immédiatement conservé sur le terrain dans du formol
DL
En laboratoire, les invertébrés, séparés des sédiments et débris végétaux par des
tamisages successifs (Reymond, 1995), sont identifiés et comptés jusqu’au niveau du
genre, de la famille ou de la classe selon les groupes (Annexe 1). La liste combinée des
DIVERSITE DU ZOOBENTHOS DANS 47 RIVIERES 11]
taxons présents chaque année dans chaque station est ensuite dressée a partir des deux
ou trois (Alpes) prélevements effectués dans chacune d’elles; cependant, le méme taxon
observé dans les deux ou trois prélèvements de la même station n’est compté qu’ une
seule fois. Toutes les analyses présentées dans cet article sont basées sur la liste
combinée des taxons présents la méme année dans la méme station. Cette approche
rend les comparaisons entre stations et entre années plus fiables en diminuant l'impact
d’evenements exceptionnels (breves crues avant le prélevement par exemple) sur la
diversité du zoobenthos (Furse et al., 1984).
Dans cette étude, la diversité des communautes d'invertébrés est estimée a partir
de huit descripteurs: (1) le nombre total de taxons (genre ou famille), (2) le nombre de
taxons sensibles aux pollutions (plécopteres, heptagéniidés et trichopteres a fourreau),
(3) le rapport nombre de taxons sensibles sur nombre total de taxons (2/1 en %), (4)
l'indice RIVAUD calculé à partir de I et 2 (Tab 1), (5) le nombre de familles, (6) le
nombre de familles sensibles aux pollutions (plécopteres, heptagéniidés et trichopteres
a fourreau), (7) le rapport nombre de familles sensibles sur nombre total de familles
(6/5 en %), (8) l'indice de diversité de Margalef (Magurran, 1988) dont l'expression est:
Diversité = (nombre de familles -1) / In du nombre d'individus récoltés.
Les descripteurs retenus ici sont ceux généralement utilisés pour l'étude des
cours d'eau (Fore er al., 1996). Certains sont très semblables entre eux (1 à 3 et 5 à 7):
tout ce qui les différencie, c'est le niveau d'identification atteint (le genre ou la famille).
Leur inclusion permet d'évaluer l’effet de ce facteur sur l’analyse des résultats.
Les diversités calculées ici ne sont pas basées sur les espèces mais sur des
genres, des familles et même des classes (Annexe 1). Cependant cette approche sim-
plifiée peut être utilisée parce qu'il existe une corrélation très significative entre le
nombre de familles et d'espèces présentes dans les rivières (Wright et al., 1998). De
plus, l'analyse des communautés d’invertebres benthiques donne souvent des résultats
analogues qu'elle soit basée sur des identifications faites au niveau au niveau de la
famille ou de l’espèce (Furse et al., 1984).
Les variations des huit descripteurs sont analysées en fonction de l'altitude des
165 stations de prélèvements, de la région où celles-ci se trouvent (Jura, Jorat et Alpes)
et de la date des campagnes de prélèvements. Trois campagnes sont comparées entre
elles: 1989-1991, 1992-1994, 1995-1997. Quatre zones d'altitude sont définies à partir
des valeurs quartiles (25 %, 50 %, 75 %) et extrêmes de l’altitude des stations de
prélèvements.
En plus de l’analyse basée sur les stations, les valeurs moyennes des huit des-
cripteurs sont calculées pour chaque rivière et pour chaque campagne de prélèvements
afin d'évaluer la tendance rivière par rivière. Pour ce faire, la valeur du coefficient de
corrélation de rang de Spearman est calculée entre les trois valeurs moyennes de chaque
descripteur et les dates correspondantes des trois campagnes de prélèvements. Le
nombre de rivières où la valeur du descripteur augmente régulièrement (r, = 1.00) est
ensuite comparée à celui des rivières où sa valeur diminue (rs = - 1.00) au moyen du
test binomial. Les rivières où les valeurs de r, sont autres que +1 ou -1 sont exclues de
la comparaison.
12 CLAUDE LANG
RESULTATS
Le tableau 2 présente l’évolution des valeurs moyennes des huit descripteurs du
zoobenthos en fonction de trois critères: |’ altitude, la région et la date des campagnes de
prélèvements. Nous constatons tout d'abord que les valeurs moyennes des huit
descripteurs augmentent significativement avec l’altitude des stations de prélèvement.
C'est entre 375 m et 449 m d'altitude, dans la zone où les impacts des activités
humaines sont les plus forts (Lang & Reymond, 1995), que les valeurs observées sont
les plus basses. Nous observons ensuite que, dans les rivières des Préalpes et des Alpes,
les nombres moyens des taxons et des familles sensibles aux pollutions sont nettement
plus élevés dans que dans celles du Jura et du Jorat. Comme la diversité totale est la
même dans les trois régions, le pourcentage de taxons et de familles sensibles aux
pollutions est plus élevé dans les rivières de montagne qu'ailleurs. Dans l'interprétation
TABLEAU 2. Valeurs moyennes par station de 8 descripteurs du zoobenthos en fonction de l'alti-
tude, de la région et de la campagne de prélèvements. Descripteurs: nombre total de taxons
d'invertébrés (NT), nombre de taxons sensibles aux pollutions (NTS), pourcentage de taxons
sensibles aux pollutions (PNTS), indice RIVAUD, nombre de familles (NF), nombre de familles
sensibles aux pollutions (NFS), pourcentage de familles sensibles aux pollutions (PNFS), indice
de diversité de Margalef (DIV). Au sein d'un même ensemble et d'une même variable, les
moyennes qui ne sont pas significativement différentes au seuil de 5 %, sont indiquées par des
lettres semblables. Ensembles 1 et 2 basés sur tous les résultats 1989 - 1997.
Ensembles Valeurs moyennes par station
comparés N° NT NTS PNTS RIVAUD NF NES PNFS DIV
1. Altitude (m) 375-449 43 18.9A 36A 172A 84A 174A 2.9A 15.6A 2.16A
450-539 43 241B 7.0B 27.9B 12.3B 20.9B 5.2B 244B 252B
340-699 48 253B 71.867 292B7 12.9B 21.755.6BE252BE2>7B
700-1410 41 26:18 104€ 40:9C 14.7C7722.0B TIC ACC
2. Région Jura) 15 7229A 60A 237A ILIA 20NA 45k 210A
Jorat®? 47 243A 64A 24.2A 11.8A 21.2A 4.8A 214A 2.53A
Alpes? 43 23.7A 10.0B 42.2B 13.9B 20.2A 6.7B 34.0B 2.62A
3. Campagne 1989-1991 165 21.7A 64A 28.0A 11.1A 19.0A 48A 244A 238A
1992-1994 165 226A 7.0A 29.1A 11.8A 19.7A 5.1A 247A 246A
1995-1997 165 26.2B 8.0B 28.9A 13.2B 22.6B 5.7B 24.4A. 2.68B
a) Nombre de stations
b) Altitude des stations: 375 - 1040 m, moyenne: 522 m
©) Altitude des stations: 430 - 775 m, moyenne: 555 m
d) Altitude des stations: 380 - 1410 m, moyenne: 882 m
des résultats, il faut noter que les effets de l’altitude et de la région sont souvent meles.
Par exemple, les valeurs moyennes observées entre 700 m et 1410 m d’altitude corres-
pondent en majeure partie aux rivières des Alpes et des Préalpes: les valeurs observées
en-dessous de 450 m, aux rivieres du Jura. Remarquons enfin que les valeurs moyennes
de six des huit descripteurs du zoobenthos augmentent entre 1989 et 1997. C'est entre la
deuxieme et la troisieme campagne de prelevements que l’augmentation est parti-
culièrement marquée.
DIVERSITE DU ZOOBENTHOS DANS 47 RIVIERES 113
22-
20 - © Press
18- ==
©)
164 ==
(tl el
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a
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N= 43 43 38 41 43 43 38 41 43 43 i 38 41
1989-1991 1992-1994 1995-1997
CAMPAGNES
FIG. 2
Variations des valeurs quartiles (petits côtés du rectangle de bas en haut: 25%, 50%, 75%) et
extremes (lignes) de l'indice RIVAUD en fonction de l’altitude des stations de prélèvements au
cours des trois campagnes effectuées entre 1989 et 1997. Les cercles et les étoiles indiquent des
valeurs aberrantes excédant 1.5 fois l’espace interquartile. Pour chaque campagne, les stations
sont réparties en 4 classes d’altitude de gauche à droite: 375 - 449 m, 450 - 539 m, 540 - 669 m,
700 - 1410 m. N = nombre de stations par classe d’altitude.
Les figures 2 et 3 montrent d’une part que les valeurs quartiles de l'indice
RIVAUD et du nombre de familles augmentent régulièrement avec l’altitude des sta-
tions dans deux des trois campagnes de prélèvements comparées, d'autre part que les
différences entre les trois campagnes sont également significatives (analyse de
variance: effet de l’altitude, P = 0.001; effets de la campagne, P = 0.001). Remarquons
cependant que, pour la campagne 1989 - 1991, les valeurs médianes de la diversité sont
les mémes dans les trois dernières zones d'altitude. Cette répartition particuliere semble
indiquer que la sécheresse exceptionnelle de 1989 a exercé une influence negative sur la
diversité du zoobenthos (Fig. 4).
La corrélation positive (r, = 0.82) qui existe entre le nombre de taxons sensibles
aux pollutions et le nombre total de taxons présents dans les 47 rivières vaudoises
étudiées entre 1989 et 1997 (Fig. 5), permet de classer celles-ci en fonction de la
114 CLAUDE LANG
40,
35 -
(O)
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1989-1991 1992-1994 1995-1997
CAMPAGNES
FIG. 3
Variations des valeurs quartiles et extrêmes du nombre de familles d'invertébrés en fonction de
l'altitude des stations de prélèvements au cours des trois campagnes effectuées entre 1989 et 1997
(légende, voir Fig. 2).
diversité du zoobenthos. En bas et à gauche de la figure, nous trouvons de petites
rivières, situées en général à basse altitude, dans des paysages très modifiés par
l’homme: la Dullive, la Petite Glâne, la Morges et le Boiron de Morges. En haut a
droite, nous observons des rivières de montagne comme l’Hongrin et la Torneresse qui
drainent des bassins versants relativement peu modifiés par l’homme, tout au moins en
amont des stations étudiées. Certaines rivières de montagne, notamment l’Avancon, la
Tinière, la Veveyse et la Grande Eau, occupent une position marginale sur la figure 5.
La diversité des taxons sensibles aux pollutions (surtout celle des plécoptères) y est très
élevée par rapport à la diversité totale. Cette situation particulière s'explique par un
caractère torrentiel accentué qui élimine de nombreux taxons. Comparons enfin la
faible diversité observée dans la Venoge à la diversité élevée de son principal affluent
le Veyron. Si la qualité de l'environnement s’améliorait à basse altitude, la situation de
la Venoge devrait se rapprocher de celle du Veyron.
L'évolution de la diversité du zoobenthos entre 1989 et 1997, pour les 47
rivières étudiées. est présentée dans le tableau 3 avec les valeurs moyennes des huit
descripteurs utilisés. Sur les 376 tendances calculées (8 descripteurs multipliés par 47
DIVERSITE DU ZOOBENTHOS DANS 47 RIVIERES 115
TABLEAU 3. Valeurs moyennes et tendances de 8 descripteurs du zoobenthos calculées pour
chacune des 47 rivieres vaudoises visitées a 3 reprises entre 1989 et 1997. Tendance: augmen-
tation (+), diminution (-), aucune tendance (0). Descripteurs: nombre total de taxons (NT),
nombre de taxons sensibles aux pollutions (NTS), pourcentages des taxons sensibles aux
pollutions (PNTS), indice RIVAUD, nombre total de familles (NF), nombre de familles sen-
sibles aux pollutions (NFS), pourcentage des familles sensibles aux pollutions (PNFS), indice
de diversité de Margalef (DIV).
Riviere Valeurs moyennes (en dessus) et tendances (en dessous)
No (Stations) NT NTS PNTS RIVAUD NF NFS = PNES DIV
1 Doye 15.3 1.7 11 DA 15.3 1.7 11 1.9
(1) 0 0 0 0 0 0 0 0
2 B. de Nyon 1557 1.8 10.6 6 15.1 1.8 11 1.9
(4) 0 0 0 0 0 0 0 0
3 Asse 19.4 3.8 16.5 8.3 17.8 3.] 15.6 DE
(4) + 0 0 + + 0 0 +
4 Promenthouse 23.8 6.7 DAD. 12 2A 4.9 22.8 2.6
(3) + + + + + + + 0
5 Colline 255 WS 28.3 13 21.8 5.5 24.7 DT]
(2) 0 + + 0 0 + + 0
6 Cordex 26.3 9.7 36.6 14.7 23.3 TI 30 2.8
@ + + + 0 0 0 0 -
7. Serine 20 6.2 30.6 11 18.7 4.8 26.3 DES
(2) 0 0 - 0 0 0 0 0
8 Dullive 10.7 0.5 I 3.5 10.7 0.5 I 1.3
(2) 0 0 0 + 0 0 0 0
9 Aubonne DD 9.3 33.9 14.5 22.9 6.2 25) 297:
(5) + 0 0 + + + 0 0
10 Toleure 26.8 10.3 38.6 14.8 22.8 Ue) 33 2.9
(2) 0 + + 0 0 0 0 -
11 Saubrette 18.3 4.7 25.3 10 LZ 4 22.9 2.1
(1) 0 0 0 0 0 0 0 0
12 B. Morges 18 2.6 13.2 ES 17.2 DIS 13.4 DAI
(5) 0 0 + 0 0 0 + 0
13 Morges 18 3 16.1 8 17.1 2.9 16.5 2
(5) 0 - - 0 0 0 0 0
14 Venoge 213 4.5 195 9.8 19 35 WZ 2.4
(14) + 0 0 + + 0 0 +
15 Veyron 30.9 10.2 32.9 15.8 24.8 TS) 29.2 5
(4) + + 0 + + 0 0 +
16 Nozon 28.9 10.8 37.1 1537 23.4 7.4 31.6 DET
(6) 0 + 0 0 0 + 0 0
17 Orbe DES 6.3 22.9 125 233 Sl DIET 2.8
(5) + + + + + + + +
18 Mujon 17.3 2 11.4 7 17.3 2. 11.4 1.9
(1) 0 0 - 0 0 0 = È
19 Arnon 30.9 10.2 32.9 15.8 24.8 18 29.2 3
(7) 0 0 + + 0 0 + 0
20 Baumine 28.7 Dall 19.3 13 25 4.7 18.3 2.8
(1) 0 0 0 0 0 0 (0) 0
21 Talent 19 4.] 18.4 8.6 11927 3.4 17.3 22
(7) + + + + + + + +
22 Buron DS 6.3 2/61 12 20.7 4.7 22.8 DIS
(2) 0 0 0 0 0 0 0 0
23 Mentue 275 8.5 30 14.1 22.9 5.8 24.5 DI]
(8) 0 0 0 0 0 0 0 0
116 CLAUDE LANG
TABLEAU 3 (suite)
Rawiere Valeurs moyennes (en dessus) et tendances (en dessous)
No (Stations) NT NTS PNTS RIVAUD "NE NFS PNES DIV
24 Sauteru 29
7.8 DAN 14.2 26 6.2 24 3). il
(2) 0 0 + 0 0 0 0 0
25 Betite:;Gläane 215.8 12 1:3 SI 15.6 122 123 2
(4) 0 + - + 0 + + 0
26 Broye 27.4 7 24.4 1331 23.4 53 22 2.8
(7) + 0 0 + + 0 0 +
27 Arbogne 23.9 5 20.4 11 21 5 18.1 DIS
(3) + + + + 0 + + 0
28 Lembe 15.7 DEN 16.5 MI 138 2.7 16.9 1.9
(1) 0 - 0 - 0 - - 0
29 Cerjaule DS 8.3 32.4 197 23 N 30.5 2.8
(1) 0 - 0 - 0 - 0 -
30 Mérine 313 EY, 37.1 16.7 IST 8 312 SA
(1) 0 0 0 0 0 0 - 0
31 Bressonne 27.8 93 32.6 14.4 23.6 6.7 28 2.9
(3) + + + + 0 + + +
32 Carrouge 21.3 4.8 21.4 10.2 19.5 4 20 22
(2) 0 0 0 0 0 0 0 -
33 Grenet 26.4 8.2 30.8 13.8 22.8 62 ZII ZA)
(4) 0 0 0 0 0 0 - +
34 Mionne 2957 10.3 34.7 1527 24.7 TR SISI DS)
(1) 0 0 0 0 0 0 + 0
35 Forestay 223 3 12.7 9 20.3 3 14.1 2.4
(1) 0 0 0 0 0 0 0 0
36 Veveyse DI. 8.7 39.3 13 18 5 doll 2.4
(1) + 0 0 + u 0 0 +
37 B.Clarens 29.4 [EZ 37.9 15.8 24.8 7.6 30.4 DIO,
(4) 0 0 0 0 + 0 0 +
38 B. Montreux 27.6 IH 40.3 1577 23.6 TE? 30.6 DIO,
(3) 0 0 0 0 0 0 - +
39 Tinière 19.7 7.9 38.8 11.6 17.4 SLI 30 DS
(3) 0 0 0 0 (0) 0 0 0
40 Grande Eau 20.8 9.5 45.7 13 17:9 6.7 STES) 2.4
(10) + - 0 0 0 0 - 0
41 Gryonne 23.8 8.9 37.6 13.6 20.8 6.1 29.6 2
(4) 0 0 - 0 0 - - +
42 P. Gryonne 25 8 32 153 225 SIT 252 ei
(1) 0 + + (0) 0 0 + +
43 Avancon 17.4 TS) 45.4 11.2 15.2 SA ST DED
(7) + + - + 0 + - +
44 Hongrin 327 1235 38.1 1783 26.7 8 30 3,2
(2) + + 0 + + + 0 +
45 Sarine 26.8 Fi] 42.2 152 223 TD 34.6 2.8
(4) + 0 0 + + 0 0 +
46 Flendruz 28 123 44.] 16.3 28 8 3922 3
(1) + + 0 + + 0 - +
47 Torneresse 30 14.1 47 17.3 24.1 8.6 355 DIO
(3) 0 0 0 0 0 0 0 *
Tendance (+/-) (17/0) (16/3) (13/5) (18/2) (3/0) Cais) (11/9) (18/5)
0
Probabilite 0 0.004 0.096 0 0.057 0.824 0.012
DIVERSITE DU ZOOBENTHOS DANS 47 RIVIERES AG
PHOSPHORE
90 -
50 -
40-
30 +
20 | T
SU T T TILE T T =
1983 1985 1987 1989 1991 1993 1995 1997
1984 1986 1988 1990 1992 1994 1996 — 1998
T = T Ter
ANNEE
FIG. 4
Evolution des concentrations moyennes en phosphore soluble (mg / m?) et des débits d'eau (mò /
sec multiplies par 10) dans la Venoge entre 1984 et 1997. Les concentrations sont pondérées par
le débit (Orand er al., 1998).
rivières), 117 indiquent une augmentation de la valeur du descripteur et 27 une
diminution (test binomial, P = 0.001); dans 232 cas, aucune tendance bien marquée
n'est observable. Lorsque les tendances sont analysées descripteur par descripteur
(colonne par colonne), les nombres d'augmentations sont significativement supérieurs à
ceux des diminutions pour six descripteurs sur huit. Pour les pourcentages de taxons ou
de familles sensibles aux pollutions, les différences ne sont pas significatives, ce qui
suggère que ces deux descripteurs sont moins efficaces que les six autres pour détecter
une tendance. Lorsqu'on examine l’évolution de la diversité rivière par rivière (ligne
par ligne), la tendance dominante est clairement visible dans la plupart des cas, même si
l'ensemble des huit descripteurs n'indiquent le même diagnostic que dans dix rivières.
Les diagnostics contradictoires (une augmentation et une diminution dans la même
rivière) sont également rares: ils sont le fait de l'indice de Margalef et du pourcentage
de taxons ou de familles sensibles aux pollutions, descripteurs dont la valeur indicatrice
a déjà été mise en doute ci-dessus.
118 CLAUDE LANG
167 |
14, o
124
105 40
TAXONS SENSIBLES
à
2- EG 12 a
10 15 20 25 30 35
TOUS LES TAXONS
Fic. 5
Relation entre le nombre total de taxons d'invertébrés et le nombre de taxons sensibles aux
pollutions (plécopteres, heptagéniidés et trichopteres à fourreau), permettant de classer les 47
rivieres vaudoises visitées entre 1989 et 1997, les unes par rapport aux autres en fonction de leur
diversité (voir Tab. 3, numéros d'identification et noms des rivieres).
DISCUSSION
Dans les rivières vaudoises, la diversité du zoobenthos augmente d’une part en
fonction de l’altitude des stations de prélèvements dans chacune des trois campagnes
effectuées entre 1989 et 1997, d’autre part en fonction du temps écoulé entre la
première et la troisième campagnes (Figs 2, 3). La première tendance correspond à
l'augmentation amont-aval des perturbations d’origine humaine qui s’observe dans la
plupart des bassins versants vaudois (Lang & Reymond 1993, 1995). La deuxième
tendance montre à la fois les progrès de l'épuration des eaux et les effets d'une météo-
rologie favorable.
L’épuration des eaux a permis de contrôler efficacement certains polluants ainsi
que l’illustre la baisse significative (r, = - 0.84, n = 14) des concentrations en phosphore
soluble dans la Venoge entre 1984 et 1997 (Fig. 4). Cette figure montre également que
la longue sécheresse de 1989 a momentanément compromis l’amélioration de la qualité
DIVERSITE DU ZOOBENTHOS DANS 47 RIVIERES 119
chimique de l’eau. En effet cette année-la, le debit moyen de la Venoge est
particulierement faible, comme d’ailleurs celui des autres rivieres vaudoises. A cause de
ce manque d’eau, les apports en phosphore soluble, ainsi que ceux d'autres polluants,
sont moins dilués et les concentrations plus élevées qui en résultent, affectent davantage
le zoobenthos (Allan, 1995).
La situation météorologique extreme de 1989 a influence fortement la premiere
des trois campagnes de prélèvements effectuées entre 1989 et 1997 (Figs 2, 3).
L'évolution ultérieure de la diversité pourrait ainsi s’interpreter comme le rétablisse-
ment progressif du zoobenthos après un épisode climatique exceptionnel. Si cette
interpretation est correcte, la diversité du zoobenthos observée pendant la dernière cam-
pagne correspondrait à une situation normale sur le plan météorologique pour les
rivières vaudoises. En prenant les diversités observées en altitude comme valeurs de
référence pour un milieu où les perturbations d’origine humaine sont minimes (Figs 2,
3), nous constatons une sérieuse baisse de la diversité à basse altitude, ceci malgré une
épuration des eaux complètement réalisée (Fiaux ef al., 1998).
L'effet de l'amélioration de la qualité chimique de l’eau sur la diversité du
zoobenthos serait ainsi contrecarré par la persistance d'autres perturbations d’origine
humaine (Allan, 1995). Citons entre autres: l’alluvionnement, l’altération des débits de
crue et d’étiage et la dégradation de la végétation riveraine. En un mot, tout ce qui
modifie le caractère naturel d'un cours d'eau et de son bassin versant fait diminuer la
diversité du zoobenthos. Le milieu naturel environnant les rivières vaudoises est
dégradé mais l'étendue de sa dégradation ne peut pas être évaluée, faute d'une base de
référence et de données quantitatives précises. Cependant, l'influence de | agriculture et
de l'urbanisation est évidente.
Les activités agricoles pratiquées dans le bassin versant influencent la diversité
du zoobenthos. L'évolution d'une rivière américaine illustre ce point (Grubaugh &
Wallace, 1995): après la disparition dans le bassin versant d'une agriculture intensive
(utilisant beaucoup d'engrais, de pesticides et provoquant une forte érosion), la
diversité du zoobenthos a fortement augmenté. Dans le canton de Vaud, l’évolution
inverse s’est produite entre 1960 et 1970 avec le passage d’une agriculture tradi-
tionnelle à une agriculture intensive (B. Reymond, com. pers.). Il est significatif de
constater que la diversité des plécoptères, insectes particulièrement sensibles à l’in-
fluence humaine (Fore er al., 1996), a fortement diminué entre 1945 et 1982 dans le
cours inférieur de la Venoge, du Talent, de la Broye et d’autres cours d’eau traversant
des paysages agricoles très modifiés par l’homme (Aubert, 1984).
En plus des effets à long terme de l’agriculture sur le paysage, des pollutions
tant aiguës que chroniques par des produits phytosanitaires pourraient expliquer la
faible diversité du zoobenthos qui persiste entre 1989 et 1997 dans certaines rivières
vaudoises où l’épuration des eaux est entièrement réalisée. Citons, par exemple, le
Boiron de Nyon, l’Asse, la Dullive, le Boiron de Morges et la Morges (Tab. 3). En
raison de la conjonction de faibles débits et d'une agriculture très active, divers produits
phytosanitaires sont présents dans l’eau en concentrations relativement élevées (Corvi
& Kim-Heang, 1997: Vioget & Strawczynski, 1997). De plus, des déversements acci-
120 CLAUDE LANG
dentels de ces produits provoquent frequemment la mort des poissons et des inver-
tebres, parfois seulement celle des invertebres; dans ce cas, les pollutions et leurs
auteurs sont particulierement difficiles a detecter (Ph. Tavel, com. pers.).
En plus de ces pollutions aigués mais sporadiques, les concentrations en phyto-
sanitaires mesurées dans ces cours d'eau suggèrent qu'il pourrait exister une pollution
chronique. Celle-ci agirait de facon subtile sur le taux de survie des especes. Par
exemple, le nombre de trichoptères du genre Limnephilus, capables d’émerger de l’eau
et donc de se reproduire, diminue significativement en présence d'une tres faible
concentration (1 ng/l) de lindane, un insecticide (Schulz & Liess, 1995). Il est bien clair
que des effets si subtils ne peuvent pas étre mis en évidence par des méthodes clas-
siques de surveillance biologique des eaux.
Le zoobenthos des rivieres vaudoises subit également les effets négatifs d’une
urbanisation accrue, surtout a basse altitude. En effet, la densité de la population
humaine qui constitue un bon indicateur des impacts lies a urbanisation (Statzner &
Sperling, 1993), augmente en général d’amont en aval dans les bassins versants du
canton de Vaud (Lang & Reymond, 1995). Une forte densité humaine entraine par
exemple: (1) l'augmentation des surfaces de sol imperméables ce qui modifie les débits
de crue et d’étiage, facteurs critiques pour le zoobenthos; (2) l’accroissement du réseau
de canalisations qui conduisent les eaux usées aux stations d’épuration (Statzner &
Sperling, 1993). Plus ce réseau est étendu, plus la matiere organique et les polluants
peuvent s’y accumuler par temps sec: de ce fait, plus grand sera impact de ces dépôts
sur la riviere apres une forte pluie, sauf si les déversoirs d’orage sont congus et entre-
tenus de facon optimale.
En conclusion, Statzner & Sperling (1993) suggèrent qu'une rivière épurée, son
bassin versant et la population qui l'habite constituent un système complexe qu'il
faudrait gérer comme un tout, en utilisant différentes approches, si l'on veut améliorer à
moindre coût la qualité biologique de l’eau. C’est une conception analogue qui devrait
être adoptée à l’avenir dans les rivières vaudoises.
REMERCIEMENTS
La collaboration d'Olivier Reymond tant sur le terrain qu’en laboratoire m’a
permis de mener à bien cette étude. Les commentaires de Michel Dethier, de Philippe
Gmür, d’Andres Strawczynski et de Philippe Vioget ont contribué à améliorer cet
article.
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122
CLAUDE LANG
ANNEXE |. Fréquence des taxons identifiés entre 1989 et 1997. Source: Lang 1996, 1997, 1998. Taxons
présents +, 1 - 10% des prélèvements, fréquents ++, 11 - 50%, très fréquents +++, 51 - 100%, - absent.
L'astérisque signale les taxons sensibles aux pollutions.
Région
Taxons®) Jura Jorat Alpes
TURBELLARIA
Planariidae
Polycelis ++ + +
Dugesiidae
Dugesia ++ ++ ++
Oligochaeta +++ +++ ar
HIRUDINEA
Glossiphonidae
Helobdella + + &
Glossiphonia + aft
Erpobdellidae
Erpobdella ++ ++ +
MOLLUSCA
Hydrobiidae + = +
Ancylidae ++ ++ x
Limnaeidae ++ + At,
Sphaeriidae ++ ++ +
Physidae + = 3
Bythinellidae + = 4
Bithyniidae = = +
HYDRACARINA +++ ++ ++
CRUSTACEA
Gammaridae +++ +++ ++
Asellidae + + +
EPHEMEROPTERA
Ephemeridae
Ephemera + + En
Heptageniidae
Epeorus* +++ +++ ++
Rithrogena* +++ ae +++
Ecdyonurus* ++ +++ +++
Heptagenia* = = n
Caenidae
Caenis + sus si
Baetidae +++ +++ +++
Ephemerellidae
Ephemerella + SEE n
Torleya - + +
Leptophlebiidae
Habrophlebia ++ ++ CE
Habroleptoides ++ ++ ++
Paraleptophlebia + + +
PLECOPTERA
Taenioptery gidae
Taeniopteryx* - < =
Brachyptera’ ++ ++ ++
Rhabdiopteryx* - + 44
Nemouridae
Amphinemura* ++ -- a
Protonemura* ++ ++ +++
Nemoura* ++ ++ ae
Leuctridae
Leuctra* +++ ++ +++
Capniidae
Capnia* + + +
Capnioneura* = z +
Chloroperlidae
Chloroperla* + = ++
Perlodidae
Isoperla* ++ en sen
Perlodes* - = +
Perlodidae gen.* - = at
a) Identifiés d'après Tachet er al. (1980). _
Sciomyzidae
Région
Taxon Jura Jorat Alpes
Perlidae
Perla* + È ann
Dinocras* - = fi
ODONATA - + =
HETEROPTERA - = n
PLANIPENNIA
Osmylidae
Osmylus # È +
MEGALOPTERA
Sialidae
Sialis + AL +
COLEOPTERA
Gyrinidae + + È
Dytiscidae + pù ni
Haliplidae ch + +
Hydrophilidae + + +
Helodidae + + A
Eubridae = + a,
Dryopididae - + &
Hydraenidae
Hydraena ++ ++ ++
Ochthebius = A a
Elmidae
Elmis ++ EME run.
Esolus ++ + +
Limnius tato +++ ++
Normandia + = È
Riolus ++ ++ +
Oulimnius + + +
Dupophilus = = A
TRICHOPTERA
Rhyacophilidae +++ +++ +++
Glossosomatidae * a + er
Hydroptilidae* ++ ++ +
Hydropsychidae +++ +++ ++
Philopotamidae - + sit
Polycentropidae + + +
Psychomiidae +++ ++ +
Phryganeidae* - = 2
Limnephilidae* +++ +++ ++
Drusinae* = = i
Goeridae* + + AL
Beraeidae* + 2 +
Odontoceridae* ++ ++ a
Sericostomatidae* Di ++ Er
Lepidostomatidae* + + Be
Leptoceridae* + = È
DIPTERA
Blephariceridae + + ++
Tipulidae ++ - Sto
Limonidae ++ +++ +++
Psychodidae ++ ++ ++
Simuliidae +++ +++ +++
Chironomidae +++ +++ it
Ceratopogonidae ++ +++ ++
Stratiomyidae + + +
Empididae +++ +++ ++
Dolicopodidae + + +
Athericidae ++ ++ +++
Dixidae = + +
Anthomyidae ~ - +
Tabanidae + + È
+ = =
REVUE SUISSE DE ZOOLOGIE, 107 (1): 123-138; mars 2000
Contribution a la connaissance de la famille Eucinetidae
(Coleoptera)
Stanislav VIT
Rue de la Poterie 26, CH-1202 Geneve, Suisse.
Contribution to the knowledge of Eucinetidae (Coleoptera). - A new
genus, Proeuzkus gen. n., is described to accomodate two new species,
P. pachys from Nepal and P. coecus from Thailand. Proeuzkus pachys
sp. n. (type-species) represents the first record of a highly specialised form
of Eucinetidae from the area extending from the Caucasus to Japan, while
P. coecus sp. n. is the first anophthalmous species of the family. Another
morphologically interesting species Tohlezkus laticanthus Sp. n. is descri-
bed from Malaysia, and Eucinetus xaca sp. n. with a highly derived aedea-
gus is added to the Mexican fauna.
Key-words: Coleoptera - Eucinetidae - Proeuzkus gen. n. - Tohlezkus -
Eucinetus - taxonomy - new species - Nepal - Thailand - Malaysıa -
Mexico.
INTRODUCTION
La famille des Eucinetidae comporte actuellement six genres et 38 especes
valides, dont 28 (plus deux sous-especes) dans le seul genre Eucinetus Germar, 1818.
Les cinq genres restants refletent une grande hétérogénéité de combinaisons des
caracteres morphologiques. Les Eucinetidae font partie des six lignées phyletiques de
Coléopteres aux comportements alimentaires corrélés avec des spores. Plusieurs
especes (larves comprises) sont connues pour étre associées a divers champignons et ıl
reste admis que certaines d’entre elles sont associées à des Myxomycetes (Lawrence &
Newton, 1980; Chandler, 1991). Les premieres formes a morphologie buccale haute-
ment spécialisée ont été décrites dans les genres Tohlezkus Vit et Jentozkus Vit (Vit,
1977), puis l’ensemble des données progressivement revu et remanié (Vit, 1981, 1985,
1990, 1995). Aucune hypothese sur les relations phylogénétiques au sein de la famille
n’a encore été publiée. En l’état de nos connaissances, on y soupçonne cependant une
grande fréquence des caractères convergeants.
Parmi des matériaux inédits, de provenances fort diverses, souvent modestes
quant au nombre d'individus disponibles, les quatre espèces réunies dans ce travail
représentent des espèces porteuses de caractères intéressants ou très particuliers. Elles
me semblent succeptibles d'apporter des éléments nouveaux au regard des relations
phylogénétiques au sein de cette famille, où des caractères vraisemblablement dérivés
se retrouvent souvent dans des groupes supposés phylétiquement éloignés.
Manuscrit accepté le 27.07.1999
124 STANISLAV VIT
Un genre nouveau, Proeuzkus gen. n. de la région orientale, est érigé pour
inclure la premiere espece hautement évoluée de la famille Eucinetidae, recensée jus-
qu'à present des vastes territoires s’etendant entre l’Iran et le Japon. Deux espèces lui
sont assignées; P. pachys sp. n., remarquable espèce du Népal (espece-type) et le minis-
cule et anophtalme P. coecus sp. n. de la Thailande. Une curieuse espece Tohlezkus
laticanthus sp. n. est décrite de Malaisie et finalement une espece du Mexique, Euci-
netus xaca Sp. n., remarquable par son édéage, complete la liste.
Abréviations utilisées:
CSV Collection S. Vit, Genève, Suisse
CUIC Cornell University, Ithaca, Etats-Unis
MHNG Museum d’histoire naturelle, Geneve, Suisse
ZML Museum of Zoology, Lund, Suede
Dans les descriptions: A.-Antennes; E.- Elytres; F.-Front; Lo.- Longueur; La. - Largeur;
m. - médian(ne); P.- Pronotum; s. - sutural(e):
DESCRIPTIONS
Proeuzkus gen. n.
Espece-type: Proeuzkus pachys sp. n.
Corps petit, en dessous de 1,5 mm, trapu, subglobuleux, rappelant celui des
Sphaerosoma (Sphaerosomatidae); téguments pigmentés, vigoureusement ponctués et
pubescents.
Tête (Fig. 1) large, encastrée dans le pronotum jusqu’au niveau des yeux, repliée
normalement sur la face ventrale du corps; yeux bien développés, latéraux: insertion
antennaire entièrement cachée au fond d’une profonde cavité, fermée ventralement par
le bord inférieur explané de la tête, et dorsalement par l'expansion latérale du front;
suture frontoclypéale fortement développée (cf. Discussion); appareil buccal (non
étudié en détail) fortement modifié, probabalement du type suceur: labre court, assez
peu sclérifié (comme chez Bisaya Reitter); palpes maxillaires quatriarticulés; palpes
labiaux triarticulés, l’article apical subulé. Antennes (Fig. 7) larges, très compactes;
formées de 11 articles fortement déprimés: scape asymétrique, pédicelle inséré sur sa
face ventrale (comme chez Jentozkus Vit). Face ventrale de la tête avec de chaque côté
un pli transversal caréné inexistant dans les autres genres, qui s’étend de la base du
submentum jusqu'aux tempes, où il forme une protubérence dentiforme saillant latéra-
lement en arrière des yeux. Pronotum enveloppant la base des élytres; scutellum
apparent. Elytres non soudés sur la suture, sans traces de stries longitudinales, mais
pourvus de 10 stries vestigiales; épipleurs entiers, faiblement élargis dans leur portion
posthumérale puis progressivement atténués, méchanisme de blocage des élytres au
repos présent.
Face ventrale (Figs 3-5). Processus prosternal bien développé: angle médian du
mésosternum très obtus, ne formant pas de processus mésosternal; cavités mésocoxales
subcontigués; métasternum grand, ses branches latérales bien développées, non
LA FAMILLE EUCINETIDAE 125
fusionnees; épisternes metasternaux libres; épisternes et épimères mésosternaux
complètement fusionnés; extrémités latérales des hanches postérieures découvertes:
cing sternites apparents chez les deux sexes; méso- et métatibias avec deux éperons
apicaux chez les deux sexes. Dimorphisme sexuel limité aux seuls épérons apicaux des
mésotibias.
Edéage (Figs 9-10) avec le lobe médian non tubuliforme et les paramères
déplacés tout à fait dorsalement rappelle celui d’ Eucilodes Vit.
Discussion. Le genre Proeuzkus est défini par son espèce-type P. pachys sp. n.
Sa tête peu modifiée, ses yeux bien développés et l’appareil buccal du type inter-
médiaire rappellent Bisaya Reitter du Lencoran et Eucilodes Vit (paléarctiques), mais
aussi Euscaphurus Casey (néarctique). Le pli transversal lamelliforme de la face ven-
trale de la tête, supportant ventralement les yeux, et terminé de chaque côté par une
saillie dentiforme est unique chez les Eucinetidae, tout comme les antennes de ce type
particulier. L’appartenance de la deuxième espèce, P. coecus Sp. n., à ce genre, est
cependant plus problématique car la tête a subi d’importantes modifications: atrophie
des yeux, évolution des pièces buccales vers le type piqueur-suceur et disparition de la
fosse antennaire (cf. sous coecus). D’autres adaptations, corrélées avec la reduction de
la taille et une biologie spécialisée, sont celles déjà connues dans d’autres genres,
comme Tohlezkus Vit ou Jentozkus Vit. Néanmoins, le recouvrement dorsal de l’inser-
tion antennaire, les antennes dilatées au niveau du funicule et déprimées, le scape
asymétrique et comprimé, la carene transversale de la tete (ici vestigiale et réduite a ses
extrémités latérales), le mésosternum court (angle médian non lancéolé) et dépourvu de
carene médiane, deux éperons apicaux des pattes, constituent les caracteres communs
aux deux espèces. P. coecus sp. n. est donc accepté ici dans le genre Proeuzkus en tant
que son espèce derivee.
Proeuzkus pachys sp. n. Figs 1, 3-11
Matériel. Holotype 4, étiqueté: NEPAL, Khandbari District, above Tashigaon, 3100 m
7.1V.1982, A. & Z. Smetana. (MHNG); Paratypes: 26, mêmes données que l’holotype, (1d
disséqué et conservé en glycérine). (MHNG); 14, 12 idem HT, “Bakan” W of Tashigaon
3200 m, 5.1V.1982, A. & Z. Smetana. (MHNG, 1% CSV).
Espece aptere chez les deux sexes, coloration d’un brun foncé rougeatre, pattes a
peine plus jaunatres; pubescence courte, couchée; téguments non translucides, brillants,
entierement densément et fortement ponctués. Corps fortement convexe, trapu, acuminé
apicalement. Longueur 1,40-1,45 mm, largeur 0,97-1,02 mm, située en avant du milieu
du corps.
Téte (Fig. 1). Large, faiblement convexe, retrécissant en arriere des yeux et
convergeant en arriere; tempes doublées ventralement, en arriere des yeux, par l’extre-
mité latérale dentiforme de la carène ventrale transversale de la tête; largeur de la tête
(yeux compris) d’un tiers supérieure à sa longueur médiane (labre non compris), cette
dernière subégale à la largeur du front. Yeux saillants latéralement et parfaitement
isolés de la face ventrale de la tête, composés d’une vingtaine de grosses ommatidies,
rebordés en bas par une fine carène subocculaire indistincte en vue dorsale. Côtés
explanés du front faiblement sinueux au-dessus de l’insertion antennaire, cette dernière
126 STANISLAV VIT
Fics 1-2
Proeuzkus spp. 1, P. pachys sp. n.: tete, vue dorsale; 2, P. coecus sp. n.: tete, vue dorsale.
Mesures données en mm.
entierement cachée au fond d’une profonde fosse antennaire entierement fermee; fron-
toclypéus court, amplement arrondi en avant, les côtés faiblement rebordes; suture
frontoclypéale fortement développée, formant un canal profond (qui semble en partie
invaginé), terminé de chaque côté par une fossette circulaire (cf. Discussion). Appareil
buccal (Fig. 6) du type spécialisé; mandibules et maxilles indistinctes; labre petit,
ramassé, peu sclérifié; dernier article des palpes maxillaires grand, étiré en pointe,
l’apex prolongé par une courte épine hyaline; palpes maxillaires triarticulés, très petits,
dépassant à peine sur le côté.
Antennes (Fig. 7) compactes, courtes (à peine aussi longues que la moitié de la
largeur basale du pronotum), fortement déprimées, très larges (seulement trois fois plus
longues que larges), leur plus grande largeur située au niveau du 8ème segment; articles
LA FAMILLE EUCINETIDAE 127
SATA WE
cr) JE
1 ANG D =
ta if
No
dì Od | WE
LG 1 = Î_, no
IERI
) Vig N 3
, 274 4
STR
Fics 3-5
Proeuzkus pachys sp. n. face ventrale du corps: 3, hanches postérieures, méta - et mésosternum
(pattes omises); 4, prosternum; 5, patte antérieure. (Mesures données en mm).
128 STANISLAV VIT
fermement encastres les uns dans les autres, portant de tres longues soies semi-dressees.
Scape en grande partie visible, ovoide, asymétrique, protubérant a son bord distal et
cachant partiellement le pédicelle; ce dernier subcylindrique, aussi long que large,
inséré sur la face ventrale du scape; articles 3-7 du funicule trés courts, fortement
transverses (au moins 3 fois plus larges que longs); articles 9 et 10 plus robustes,
transverses, deux fois plus larges que longs, (évoquant une massue mal différenciée);
segment apical subovoide, plus large que long, surmonté d’une pubescence apicale.
Pronotum tres grand, occupant un quart de la longueur totale de l’animal. Base
doublement sinueuse mais peu projetée en arriere; scutellum grand, triangulaire, entie-
rement visible: côtés finement rebordés; angles postérieurs aigus, enveloppant la base
des élytres. Ponctuation forte, plus espacée que celle de la tete ou celle des élytres.
Elytres sub-hémisphériques, aussi longs que larges ensemble; côtés fortement
arqués; suture élytrale non soudée; strie juxtasuturale présente, raccourcie, marquée tout
au plus dans la moitié apicale; interstrie juxtasutural renflé dans le tiers apical.
Ponctuation grossiere et dense, non alignée en stries longitudinales; dix stries ponctuées
demeurent cependant nettement distinctes sur chaque élytre en milieu aqueux, les stries
2 et 4 raccourcies, ne dépassant pas le tiers basal.
Face ventrale (Figs 3-5) brillante et aussi fortement ponctuée que la face dorsale,
aire médiane faiblement déprimée. Epipleures entiers, faiblement élargis dans leur
portion posthumérale, puis longuement atténués. Limite médiane du mésosternum
simplement obtuse, angle apical renflé, glabre, non étiré en un processu médian sépa-
rant les hanches: métasternum grand, sutures transverses et suture médiane indistinctes;
languette métasternale bien développée, courte, subparallèle; metépisternes trapus, sub-
triangulaires (comme chez Eucinetus apterus Vit, 1990), le bord externe explané,
formant un dispositif de blocage des élytres; mesépimeres transverses, fusionnés avec
les mesépisternes, la suture indistincte, leur structure tégumentaire cependant diffé-
renciée. Plaques coxales larges, mais orientées longitudinalement, laissant les extré-
mités latérales glabres des hanches postérieures largement découvertes: bords latéraux
rebordés, arqués puis discrètement sinués avant les angles apico-externes; ces derniers
presque droits, émoussés, peu saillants en arrière, n’atteignant cependant pas le bord
postérieur du premier segment abdominal: bords postérieurs faiblement concaves et
pourvus d’une nette encoche proche de la ligne médiane; premier segment abdominal
pourvu d’une courte carène médiane: le 5ème rebordé; sternites 6 et 7 faiblement
sclérifiés, rétractés au repos à l’intérieur de l'abdomen.
Pattes (tarses compris) peu modifiées, les tibias subcylindriques, s’élargissant
progressivement vers l’apex. Fémur de la patte antérieure (Fig. 5) simple, plus long que
le tibia, ce dernier plus long que le tarse; protarses sans dimorphisme sexuel: les deux
éperons apicaux du protibia petits, pratiquement indistincts; mésotibia faiblement
recourbé, armé sur sa face externe de nombreuses (12-15) épines plus fortement pig-
mentées; apex avec deux éperons de longueur subégale chez les deux sexes; éperon
externe fortement modifié chez le mâle (Fig. 8); métatibia coudé à la base, aussi long
que le tarse, face externe couverte de 12-15 épines effilées et plus fortement pigmen-
tées, non scindées en deux groupes distincts; apex avec deux éperons dont le plus grand
LA FAMILLE EUCINETIDAE 129
atteint la base du peigne apical du segment basal du tarse; ce dernier subégal aux
articles 2, 3, 4 ou 3, 4 et 5 réunis.
Edéage (Figs 9, 10) avec le lobe médian tres large, simplement lamelliforme,
dépourvu de renflure apicale; paramères amplement arrondis apicalement, très
rapprochés dorsalement. Ce type d’edeage présente quelques analogies avec celui
d’Eucilodes Vit (Vit, 1977, 1985).
Biologie inconnue, mais il s’agit d’une espèce récoltée au-dessus de 3000 m.
Distribution. Connu seulement de la localité-type: Népal, Kandbari district.
Discussion. Le caractère externe le plus frappant de P. pachys sp. n. est l’aspect
du tégument: épais, brillant, il est vigoureusement ponctué et exempt de microréti-
culations et de ponctuation nettement rapeuse. Or la ponctuation franche et grosse n'est
connue que chez de grandes espèces ailées (Eucinetus Germar, Nycteus Latreille,
Eucilodes Vit, Euscaphurus Casey), où elle est confinée à la face dorsale de la tête.
Chez P. pachys elle s’étend, presque identique, sur l’ensemble de la face dorsale et
ventrale du corps.
Quant a la conformation de la suture frontoclypéale, formant une sorte de
gouttiere invaginée, flanquée de chaque coté d’une fossette, elle demeure également
unique. Sa présence est d’autant plus surprenante qu’on releve, chez les Eucinetidae,
une tendance certaine a la disparition de la suture frontoclypéale chez toutes les petites
especes (y compris Bisaya nossidiiformis Reitter) présentant un appareil buccal du type
modifié. I] pourrait s’agir d’un dispositif rudimenraire du type mycangium (Crowson,
1981), destiné au transport des spores de champignons (notamment Myxomycetes),
dont des Eucinetidae sont supposés se nourrir. Cependant des données attestees,
concernant les particularités alimentaires des Eucinetidae, restent encore trop pauvres
(Lawrence & Newton, 1980; Chandler, 1991).
Proeuzkus coecus sp. n. Fig. 2
Matériel. Holotype 9%, étiqueté: Thailand Doi Inthanon 1750 m. 7.X1.85 Löbl,
Burckhardt. (MHNG); Paratype. 1 ®, étiqueté: /Thailand-Chiang Mai Doi Saket “1130 m. 4.XII.
87” P. Schwendinger. (MHNG - exemplaire disséqué conservé en glycérine).
Wie]
Espece aptere et anophtalme (femelle), coloration entierement d’un jaune clair
translucide, épines latérales et cils apicaux des tibias et des tarses fortement rembrunis,
noiratres; pubescence des téguments fine, couchée. Corps trapu, une fois et demie plus
longue que large, régulièrement convexe, peu atténué apicalement. Longueur 0,86 mm,
largeur 0,55 mm, située au niveau posthumeral.
Téte (Fig. 2). Large (une fois et demie plus large que longue sur la ligne mé-
diane, labre non compris), modérément convexe, paralléle au niveau des tempes et
presque semi-circulaire en avant; tempes doublées ventralement, en dessous de l’inser-
tion antennaire, par une lame latérale dentiforme non prolongée ventralement sous
forme d’une carène transversale de la tête. Yeux absents; côtés du front fortement
explanés en avant de l’insertion antennaire; fosses antennaires nulles car non fermées
ventralement par le bord inférieur explané de la tête, développé chez d’autres espèces.
Appareil buccal du type piqueur-suceur; suture frontoclypéale absente; clypéus ample-
ment arrondi, échancré au milieu, finement rebordé; ponctuation forte, éparse, granu-
leuse.
130 STANISLAV VIT
Antennes compactes, deprimees, courtes (plus courtes que la moitié de la largeur
basale du pronotum), larges (seulement quatre fois plus longues que larges), leur plus
grande largeur située a peu pres au niveau du Seme segment; articles fermement
encastres les uns aux autres. Scape visible dorsalement, fortement comprimé, étroit,
allongé, et protuberant à son bord distal, cachant pariellement le pédicelle; ce dernier
robuste, subcylindrique,inséré sur la face ventrale du scape; articles 3 a 7 du funicule
fortement transverses (deux a trois fois plus larges que longs); articles 8 a 10 plus
allongés, évoquant une massue mal différenciée: segment apical réduit, subovoide, plus
long que large, surmonte d’une pubescence apicale.
Base du pronotum doublement sinueuse, fortement projetee en arriere, suplom-
bant celle des éiytres; scutellum grand mais entièrement caché sous le rebord basal du
pronotum; côtés droits, finement rebordés; angles postérieurs aïgus, étirés en arrière.
Ponctuation fine et éparse sur le disque, plus grossière dans les déclivités latérales.
Elytres d’un tiers plus longs que larges ensemble: côtés arqués, acuminés vers
l’apex; suture élytrale non soudée; strie juxtasuturale et stries longitudinales absentes;
six stries vestigiales, rudimentaires et incomplètes demeurent néanmoins perceptibles
lors d’un examen de l’élytre dans une solution aqueuse.
Face ventrale avec une forte ponctuation serrée sur les plaques coxales et le
metasternum; aire médiane des plaques coxales à peine déprimée.
Epipleures entiers, étroits, atténués à l’apex. Hanches médianes contigües; limite
médiane du mésosternum obtuse, angle apical renflé, glabre, non étiré en un processus
médian séparant les hanches: métasternum petit; languette médiane longuement effilée,
se prolongeant jusqu’au bord postérieur des plaques coxales sous forme d’une fine
carène médiane: branches latérales étroites, entièrement fusionnées avec les métépis-
ternes; dispositif de blocage des élytres présent: mésépimères fusionnés avec les
mésépisternes. Plaques coxales larges mais orientées longitudinalement, laissant décou-
vertes les extrémités latérales glabres des hanches postérieures: bords latéraux délica-
tement rebordés, faiblement arqués: angles apico-externes aigus, projetés en arrière, au-
dessus du bord postérieur du premier segment abdominal; bords postérieurs doublement
sinueux. Cinq sternites visibles chez la femelle, le cinquième large, ogival, très fine-
ment rebordé, aussi long que les sternites 3 et 4 réunis.
Pattes (tarses compris) très robustes, les tibias déprimés, s’élargissant fortement
vers l’apex. Protibia aussi long que le tarse, apex sans éperons apicaux distincts; fémurs
des pattes antérieures et médianes avec une saillie dentiforme contigüe au trochanter;
face externe du mésotibia avec deux rangées longitudinales, serrées, de soies spini-
formes plus pigmentées: apex avec deux éperons effiles; métatibia aussi long que le
tarse, face externe du metatibia garni de nombreuses soies spiniformes rembrunies, for-
mant un groupe préapical plus ou moins nettement séparé, apex avec deux éperons
effilés plus longs que le segment basal du tarse; ce dernier plus court que les articles 2,
3, 4 ou 3, 4 et 5 réunis.
Distribution. Les deux captures proviennent de la province de Chiang Mai au
nord de la Thailande, de tamisages de débris végétaux en forét de montagne, entre 1200
et 1700 m.
LA FAMILLE EUCINETIDAE 131
Fics 6-11
Proeuzkus pachys sp. n. : 6, appareil buccal, vue ventrale; 7, antenne; 8 mésotibia et les éperons
apicaux (d); 9, édéage, vue ventrale; 10, édéage, vue latérale; 11, segments abdominaux VI et
VII (3). (Mesures données en mm).
132 STANISLAV VIT
Discussion. L’ anophtalmie de cette espèce et la présence d’un puissant appareil
buccal du type piqueur-suceur ont pour corollaire de tres notables modifications dans la
morphologie de la tête: de longues tempes parallèles: bords latéraux du front très
fortement explanés; échancrure gulaire occupant pratiquement toute la face ventrale de
la tête, d’où la suppression du pli transverse de la tête, dont il ne subsiste que les
extrémités latérales dentiformes: absence du bord latéral inférieur de la tête qui
normalement constitue la paroi ventrale de la fosse antennaire. D’autres modifications à
caractère convergent, que l’on retrouve chez d’autres espèces et genres de la famille:
fusion des pièces sternales, absence de la strie juxtasuturale, réduction des stries
élytrales ou réduction du nombre de sternites apparents, sont à mettre en corrélation
avec la réduction notable du corps (0,86 mm = la plus petite espèce connue). Il serait
utile de pouvoir connaître le mâle et la structure de l’édéage de cette espèce.
Tohlezkus laticanthus sp. n. Figs 12-16
Matériel. Holotype d , étiqueté: MALAYSIA: Pahang, 4 mi NE Cameron Highlands, 23-
25.IV. 1977, (berlese leaf litter), L. Watrous. (CUIC).
Espece aptere (pas d’ailes distinctes par transparence des téguments chez le
male), coloration entièrement d’un brun roussatre clair, côtés de la tête rembrunnis, cils
apicaux des tibias et des tarses a peine plus foncés: pubescence fine, couchée. Corps
allongé, deux fois plus long que large, faiblement acuminé apicalement. Longueur 1,05
mm, largeur 0,62 mm, située au niveau posthumeral.
Tete. Large, subogivale, modérement convexe, parallele au niveau des tempes;
côtés convergeant en avant; clypéus arrondi; ponctuation forte, subräpeuse. Yeux bien
développés, latéraux, formés d’un gros grain hyalin fortement saillant latéralement;
limites latérales du cranium fortement pigmentées; insertion antennaire entierement
cachée sous le bord explané du front; cavité antennaire fermée ventralement. Appareil
buccal du type piqueur-suceur; suture frontoclypéale absente; bord antérieur reborde.
Antennes compactes, peu déprimées, un peu plus longues que la moitié de la
largeur basale du pronotum, épaisses, au moins 5 fois plus longues que larges, leur plus
grande largeur située au niveau du 8ème segment; articles encastrés les uns dans les
autres. Scape en partie caché sous le rebord explané du front, cylindrique; pédicelle
cylindrique, subcarré, inséré axiallement; articles 3 à 6 du funicule courts, transverses
(deux fois plus larges que longs); articles 7 a 11 plus robustes, plus allongés, faiblement
transverses; segment apical grand, subovalaire, plus long que large, surmonté de
pubescence apicale.
Base du pronotum doublement sinueuse, projetée en arrière, suplombant la base
des élytres; scutellum petit, entièrement caché: côtés très finement rebordes; angles
postérieurs aigus, étirés en arrière. Ponctuation nette sur le disque, plus grossière dans
les déclivités latérales.
Elytres une fois et demie plus longs que larges ensemble, ratio Lo.s./La. = 1,46:
côtés peu arques, subparallèles, faiblement acuminés; vraisemblablement soudés sur la
suture; strie juxtasuturale oblitérée; stries longitudinales absentes: cinq stries ponctuées
vestigiales et incomplètes cependant perceptibles dans la portion basale lors d’un
LA FAMILLE EUCINETIDAE
Fics 12-15
Tohlezkus laticanthus sp. n.: 12, édéage, vue ventrale; 13, segments abdominaux VI et VII (£):
14, hanche antérieure et profémur (normalement constitués): 15, patte antérieure gauche (térato-
logique). Mesures données en mm.
134 STANISLAV VIT
examen dans une solution aqueuse. Ponctuation tres serrée et subräpeuse, arrangee, pres
de la base, partiellement, en paliers transverses .
Face ventrale entierement, densement, et räpeusement ponctuée; aire médiane
des plaques coxales nettement déprimée.
Epipleures entiers mais de forme particulière: d’abord étroits et fortement
comprimés dans leur portion post-humérale, puis larges, s’élargissant même progressi-
vement jusqu'au niveau du troisième sternite et brièvement atténués vers les angles
suturaux. Processus mésosternal semblable à celui rencontré par exemple chez Eusca-
phurus spinipes Vit; élancé, cordiforme, pourvu d’une carène médiane, acuminé apica-
lement, mais ne séparant pas les hanches médianes; cavités cotyloïdes contigués.
Métasternum petit; languette médiane fortement éffilée, se prolongeant jusqu’au bord
postérieur des plaques coxales sous forme d’une fine carène médiane; branches latérales
étroites, tout au plus faiblement élargies à leurs extrémités latérales; entièrement
fusionnés avec les métépisternes; mésépimères, fortement transverses, subfusionnés,
leur suture avec le mesépisterne distincte: mésosternum et mésépisternes fusionnés.
Plaques coxales orientées longitudinalement, étroites, laissant largement découvertes
les extrémités latérales glabres des hanches postérieures: bords latéraux délicatement
rebordés, faiblement arqués; angles apico-externes aigus, émoussés au sommet, forte-
ment projetés en arrière au-delà du bord postérieur du premier sternite; bords posté-
rieurs fortement convergents vers la ligne médiane. Cinq sternites visibles ( sûrement
chez les deux sexes), le cinquième large, ogival, finement rebordé, aussi long que les
sternites 3 et 4 réunis. Chez le mâle les sternites 6 et 7 (Fig. 13) restent faiblement
sclérifiés et entièrement retractés dans l'abdomen.
Pattes (tarses compris) robustes, déprimées, tibias s’elargissant fortement vers
l’apex; profémur (Fig. 14) avec une saillie dentiforme contigüe au trochanter et un
peigne de soies longeant le bord externe: patte tératologique (cf. Remarque) de cette
exemplaire (Fig. 15) présente un éperon apical et le protarse non dilatés chez le mâle
(!); mésotibia faiblement incurvé, plus court que le tarse; face externe avec deux ran-
gées longitudinales serrées de soies spiniformes, rembrunies, apex avec deux éperons
dont l’interne fortement développé chez le mâle, épais, incurvé et renflé apicalement,
presque aussi long que le segment basal du tarse: métatibia faiblement incurvé, aussi
long que le tarse; face externe garnie de nombreuses soies spiniformes plus pigmentées
et d’un peigne préapical d’épines foncées, apex avec deux éperons dont un remar-
quablement développé, en forme de “chausse-pied” (Fig. 16), dilaté, spatuliforme,
tronqué apicalement, plus long que le segment basal du tarse; ce dernier subégal aux
segments 2, 3, 4 ou 3, 4 et 5 réunis.
Edeage (Fig.12) pourvu de structures membraneuses internes et de parameres
explanés du côté dorsal et finement ciliés a l’apex.
Caracteres sexuels. Un seul male étant connu, il n’est pas possible de définir si
l’éperon apical modifié des métatibia (Fig.16) est, ou n’est pas, un caractère sexuel du
mâle (au même titre que celui des mesotibias), ou un caractère propre de cette espèce.
Remarque. Les pattes antérieures de cet exemplaire unique sont incomplètes
(Figs 14-15). A l’une manque le tibia, l’autre est sujette à une malformation curieuse,
où son tibia, de surcroît malformé, est articulé directement avec la hanche.
LA FAMILLE EUCINETIDAE 135
Discussion. Les deux caractères remarquables de cette espèce sont ses épi-
pleures dilatés dans la partie subapicale et I’hypertrophie de l’éperon apical des méta-
tibias. On relève néanmoins, ou niveau de la famille, plutôt une tendance à la réduction
des épipleurs et une régression des éperons apicaux des tibias.
Eucinetus xaca Sp. n. Figs 17-20
Matériel. Holotype d , étiqueté: Mexico: Oaxaca, 12 km N Oaxaca City, hwy 175 1900
m. 28.IX. 1990, sifting litter at small stream, tropical mountane forest. (ZML). Paratypes. 22 9
mêmes données que l’holotype; 1d (1.X. 1990), 29 © (5.X. et 8.X.1990) Mexico: Oaxaca, 12
km N Oaxaca City, hwy 175 1900 m. 1.-8.X.1990 leg. R. Raranowski, Pit-fall traps near small
stream, trop. mount. forest. (ZML); 25 4, 29 2 Mexico: Oaxaca 12 km N Oaxaca 2000 m.,
15.IX.1986 R. Baranowsk, sifting litter in creek, trop. mont. forest; 26 4,29 © Mexico: Oaxaca
10 km. N Oaxaca 1900 m. 13.1X. 1986 R. Baranowski, sifting litter at small stream, trop.mont.
forest. (ZML, CSV). Autres. 1% (xaca ?) Jalisco 50 km S Puerto Vallarta 800 m. 16.V1.1986 leg.
M. Sörensson & B. Martensson (ZML).
Longueur dorsale médiane (tête non comprise) 3,1-3,6 mm, corps subparallele,
la plus grande largeur 1,73-1,86 mm; ratio Lo.m. (sans téte)/ La. = 1,97-2,01 Coloration
presque uniforme, d’un brun rougeätre foncé, brillant, partie apicale des élytres parfois
sombre: pattes plus claires: antennes tricolores, claires et noirätres.
Tête. Transverse, longueur médiane inférieure à la largeur du front entre les
yeux, ratio Lo.m./La.F. = 0,83-0,92; ponctuation fine, serrée.
Antennes tricolores (articles 1-3 bruns, 4-9 noirätres, 10 et Il jaunätres),
longues environ comme la base du pronotum mesurée en vue dorsale, ratio Lo.A./La.P.
= 0,95-1,11; tous les articles allongés; pédicelle trois fois plus long que large, plus long
que le 3ème; ce dernier grêle, tout juste plus court que le 4ème; articles 4 à 10
subégaux, diminuant progressivement en longueur; | 1ème article une fois et demie plus
long que large.
Ponctuation du pronotum fine et serrée sur le disque, aussi forte que celle de la
tête dans les déclivités latérales.
Elytres longuement subparallèles, ratio Lo.m./La.(sexes confondus) = 1,1,60-
1,74, rebord latéral perceptible de dessus au moins dans le tiers apical. Strie juxta-
suturale oblitérée a environ trois longueurs de scutellum de la base; interstrie juxta-
suturale faiblement enflé à l’apex; stries longitudinales seulement peu visibles, mar-
quées tout au plus dans la moitié apicale, interstries à peine convexes; ponctuation fine,
paliers transverses très serrés (4 paliers = 0,1 mm).
Face ventrale. Epipleures raccourcis, se confondant avec le bord de l’élytre au
niveau du 4e sternite; leur largeur après l’étranglement posthuméral égale à la moitié de
la longueur du 3ème article antennaire. Metasternum densement ponctué; languette
métasternale longue, subtriangulaire, suture médiane nettement imprimée; sutures trans-
verses distinctes. Mesurés sur leur bords externes, les métépisternes sont aussi longs
que les mésépimères, ces derniers presque aussi larges que longs; suture séparant les
mésépimères des mésépisternes nette, non renflée. Plaques coxales larges, transversales,
cachant parfaitement les extrémités latérales des hanches postérieures; côtés non
rebordés: angles apico-externes arrondis (atteignant le niveau du bord postérieur du
premier segment abdominal); bords postérieurs simples, convergeant vers la ligne
médiane. Bord postérieur du Se et 6e sternite amplement échancré (à ).
186 STANISLAV VIT
TI RER
RR a
TTS!
/
SS
pri
PAZZO
urn! Zu
Stino
Fics 16-20
Tohlezkus laticanthus sp. n.: 16, éperon apical des métatibias; Eucinetus xaca sp. n.; 17, édéage,
vue ventrale; 18, édéage, détail de la partie apicale du lobe médian: 19, édéage, vue latéral du
méme; 20, patte postérieure, tibia et tarse. Mesures données en mm.
LA FAMILLE EUCINETIDAE 137
Patte antérieure.Tarse fortement dilaté chez le male; subégal chez les deux sexes
à la longueur du tibia, ce dernier modérément renflé chez le mâle, subparallele chez la
femelle.
Patte médiane. Tibia très nettement plus court que le tarse, progressivement
élargi, bord externe avec 3 à 8 épines noires, couchées, apex avec un seul éperon peu
développé chez la femelle, deux éperons de longueur subégale chez le mâle, l’éperon
interne simplement épaissi.
Patte postérieure (Fig. 20). Tibia nettement plus court que le tarse, légèrement
incurvé, garni sur sa face externe de 13 - 20 épines semi-dressées, plus 2 - 4 épines
dans le groupe pré-apical pas nettement singularisé; un seul éperon apical, aussi long
que la moitié du basitatarse; longueur de ce dernier nettement inférieure à celle des
articles 2, 3, 4 ou 3, 4 et 5 réunis.
Edéage (Figs 17-19) pourvu de puissantes excroissances dentiformes à la base
de l’étranglement subapical du lobe médian, portion distale des paramères lamelliforme
(cf. Discussion).
Distribution. Mexique méridional, Oaxaca.
Discussion. Avec d'importantes modifications de la portion apicale des para-
mères (normalement arrondie) et de la portion subapicale du lobe médian (normalement
tubuliforme), muni ici de puissantes excroissances latérales fortement sclérifiées,
l’édéage de Eucinetus xaca sp. n. confirme une évolution morphologique tout a fait
interessante. Ce type d’édéage a déjà été signalé (Vit, 1990) pour deux autres espèces
du groupe haemorrhoidalis, à savoir: E. pecki Vit et E. apterus Vit, décrites également
du Mexique. Or cette évolution vers une forme différente de l’édéage - pour l'instant
parfaitement isolée au sein du genre Eucinetus - apparaît chez des espèces mexicaines
de façon tout à fait endémique. Elle ne se rencontre ni au nord, chez les espèces néarc-
tiques, ni au sud, chez celles néotropicales. L’édéage d’E. xaca Sp. n. est très proche du
celui d’E. apterus, l'unique espèce aptere du groupe haemorrhoidalis, bien caractérisée
par ailleurs par une réduction des métépisternes, relative à la disparition des ailes
métathoraciques.
REMERCIEMENTS
Je tiens à remercier ici Dr C. Besuchet et Dr I. Löbl du Muséum d'Histoire
naturelle de Genève, Dr A. Smetana d'Ottawa, Dr Quentin D. Wheeler du Département
d’entomologie de l’Université Cornell d’Ithaca et Dr R. Danielson du Muséum de
Zoologie de Lund, pour m'avoir confié l’étude des matériaux traités dans cette note.
BIBLIOGRAPHIE
CROWSON, R. A. 1981. The Biology of the Coleoptera. Academic Press, London, 802 pp.
CHANDLER, D. S. 1991. Comparison of some slime-mold and fungus feeding beetles (Coleoptera:
Eucinetoidea, Cucujidoidea) in an old-growth and 40-year-old forest in New Hampshire.
The Coleopterists Bulletin 45(3): 239-256.
LAWRENCE J. F. & NEWTON A. F. Jr. 1980. Coleoptera associated with the fruiting bodies of
slime molds (Myxomycetes). The Coleopterists Bulletin 34(2): 129-143.
138 STANISLAV VIT
Vit, S. 1977. Contribution à la connaissance des Eucinetidae (Coleoptera). Revue suisse de
Zoologie 84: 917-935.
VIT, S. 1981. Une nouvelle espèce du genre Tohlezkus Vit de Taiwan (Coleoptera, Eucinetidae).
Revue suisse de Zoologie 88: 769-774.
Vir, S. 1985. Etude de la morphologie des espèces paléarctiques du genre Eucinetus Germar et
quelques remarques sur son utilisation taxonomique (Coleoptera: Eucinetidae). Revue
suisse de Zoologie 92: 421-460.
Vit, S. 1990. Revision des espèces néotropicales du genre Eucinetus Germar (Coleoptera: Euci-
netidae). Naturaliste canadien (Revue d’Ecologie et de Systématique) 117: 103-122.
Vit, S. 1995. Deux especes nouvelles d’Eucinetidae d’ Amérique du Nord particulierement inte-
ressantes (Coleoptera: Eucinetidae). Elytron 9: 125-137.
REVUE SUISSE DE ZOOLOGIE 107 (1): 139-151; mars 2000
Spatio-temporal distribution of size classes and larval instars of
aquatic insects (Ephemeroptera, Trichoptera and Lepidoptera) in a
Potamogeton pectinatus L. bed (Lake Geneva, Switzerland)
Ruth BÄNZIGER
Laboratoire d'écologie et de biologie aquatique, 18 ch. des Clochettes,
University of Geneva, CH-1206 Geneva, Switzerland.
Spatio-temporal distribution of size classes and larval instars of aquatic
insects in a Potamogeton pectinatus L. bed (Lake Geneva, Switzerland). -
Temporal changes of aquatic insect instars or size classes were monitored ın
different parts of a Potamogeton pectinatus bed. The hypothesis of a diffe-
rent distribution of the aquatic insects in the macrophyte bed according to
their life stage and of a spatio-temporal segregation of congeneric species
was tested. Head capsules widths of seven insect species (Caenis horaria
and C. luctuosa (Ephemeroptera), Mystacides azurea, M. longicornis,
Oecetis lacustris and O. ochracea (Trichoptera) and Acentria ephemerella
(Lepidoptera)) were measured at monthly intervals from May to November
1994. Samples were taken in the edge and in the centre of the macrophyte
bed on all sampling occasions, and in May, June and July, additional samples
were taken from the 2 m sediment belt adjacent to the macrophyte bed. Each
couple of congeneric species showed segregation by size before hibernation
and showed delayed emergence patterns.
Key-words: Lake - Ephemeroptera - Trichoptera - Lepidoptera - macro-
phyte - distribution - size class.
INTRODUCTION
Habitat segregation or space partitioning among closely related species have
been often addressed in rivers (Malas & Wallace, 1977; McAuliffe, 1984). Few papers,
however, have been concerned with this subject in lakes. Species with similar
ecological niches are not always separated in space and time, thus competition is not
necessarily involved and some congeneric species do coexist (Hildrew & Edington,
1979). Segregation, if there is any, is therefore likely to occur at another scale than at
the species level. In larval insects or nymphs, delayed growth of the last instars or
nymphal stages have been observed, often resulting in temporal segregation of
emergence (Macan, 1965; Tudorancea & Green, 1975; Malas & Wallace, 1977;
Bengtsson, 1981; Sweeney & Vannote, 1981; Brittain, 1982). Hildrew & Edington
(1979) showed that two congeneric hydropsychid caddisflies avoided coexistence by
different microhabitat colonization of some instars. Hydropsychidae were also studied
Manuscript accepted 13.09.1999
140 RUTH BÄNZIGER
by Muotka (1990) who showed them having different microhabitat preferences accor-
ding to different larval stages. Minshall (1984) observed changes in spatial distribution
of insects according to developmental stages. Altogether, between egg and adult, the
growing insects may live through many different feeding modes, behaviour or habitat
niches (Winterbourn, 1971; Resh, 1979; Palmer et al., 1993). These changes according
to development allow them to reduce competition if resources are scarce.
In a previous paper (Bänziger er al., subm.), we demonstrated differences in
densities of several invertebrate taxa between the edge and the centre of macrophyte
beds. The question arose whether these differences in density could be related to larval
instar or size class distribution.
The purpose of the present study was to test the following hypotheses in a
Potamogeton pectinatus L. bed of the littoral zone of Lake Geneva: 1) distribution of
insects in macrophyte beds changes according to larval instar or size class; and ii)
congeneric species living in the same macrophyte bed differ in size and/or timing of
larval instars.
MATERIAL AND METHODS
The samples were taken in a Potamogeton pectinatus L. bed near Corsier
(46°16' N, 6°12' E) in the littoral zone of Lake Geneva, Switzerland.
Samples were taken by scuba diving at a depth of 3.0 - 3.5 m in the edge, in the
centre and adjacent to the macrophyte bed. The edge was defined as the margin of the
bed characterised by lower macrophyte stem density, the presence of more filamentous
algae (depending on the season) and by shorter shoots than in the centre of the bed.
This corresponded to the 2 m wide outer belt of the macrophyte bed. Macrophyte
density in the centre of the bed was 80 stems m’? at maximum density, i.e. in July. The
(arbitrarily) 2 m wide area around the macrophyte bed, consisting of sediment, was
defined as the adjacent sediment.
Macrophyte samples were taken from May through to November 1994, adjacent
sediment was sampled from May to July. When the macrophytes senesced - from
October to November - sampling was carried on in the plant underlying sediments and
the remains of the Potamogeton bed. Collections were made using different sampling
gears depending on the substrate:
Collections of fully grown macrophytes were made using a sampler modified
after Gerking (1957) with a surface area of 0.25 m? and a height of 1 m. When
vegetation was less dense a frame of 0.0625 m? with an attached net was used. Each of
the two sampling gears was lifted down on the macrophytes by a scuba diver, the plants
were teamed out of the sediment and the sampler was closed: the Gerking-like sampler
by a trap and the net by a string.
Sediments underlying the macrophyte bed were collected using corers covering
an area of 0.005 m? and pushed 10 cm deep into the sediments. Adjacent sediments
were collected inside a 0.25 m? frame using the same net as for vegetation samples.
In the laboratory, samples were thoroughly rinsed with tap water and the
macroinvertebrates were retained in a 250 um sieve. They were conserved in 4%
formalin.
AQUATIC INSECTS IN MACROPHYTE BEDS 141
Seven insect species were retained for measurements: Caenis horaria (L.), C.
luctuosa (Burm.) (Ephemeroptera), Mystacides azurea (L.), M. longicornis (L.), Oecetis
lacustris (Pictet), O. ochracea (Curtis) (Trichoptera) and Acentria ephemerella (Denis
and Schiffermiill.) (Lepidoptera) on the basis of their abundance in the samples, their
identification and/or the availability of congeneric species.
Larvae of mayflies were assigned to size classes as they do not have easily
distinguishable cohorts (Benke & Jacobi, 1986). The size classes of Caenis spp. and the
instars of Trichoptera and Lepidoptera were assigned on the basis of head capsule
widths (HCW) as they were often reported to be more reliable than body length in
separating the different stages (Bradbeer & Savage, 1980; Bass ef al., 1982).
Measures of HCWs were made at their widest point including the eyes (Smock,
1980; Bass er al., 1982). All measures were made with an accuracy of 0.025 mm using
a dissecting microscope with a micrometer.
Head capsules widths of Caenis spp. were divided into 18 classes of 0.05 mm
each (Table 1). Larvae of Mystacides spp. and Oecetis spp. were divided into five larval
instars (Table 2). At instar I, genera could not be keyed to species, so they were termed
“juveniles”.
TABLE |
Correspondance between size classes and head capsule widhts (HCW) of Caenis spp.
size classes HCW of Caenis size classes HCW of Caenis size classes HCW of Caenis
spp. (mm) spp. (mm) spp. (mm)
I 0.125 - 0.2 U 0.575 - 0.65 13 1.025 - 1.1
2 0.2 = 0.275 8 0.65 - 0.725 14 ILES
3 0.275:- 0.35 9 0.725 - 0.8 15 12.1779%-.1:25
+ 0.35 - 0.425 10 0.8 - 0.875 16 E25 les 25)
5 0.425 - 0.5 11 0.875 - 0.95 17 1.325 - 1.4
6 12
050575 2 0:9377.1:025 18 1.4 - 1.45
Data in the literature dealing with HCWs of A. ephemerella were scarce. Five
instars were reported in the literature and Haenni (1974, 1980) identified the sizes of
the different instars by collecting individuals in the field and by rearing them.
TABLE 2
Correspondance between instars and head capsule widhts of Mystacides azurea, M. longicornis,
Oecetis lacustris and O. ochracea.
instars HCW of instars HCWofM. instars HCW of instars HCW of O.
Mystacides longicornis Oecetis lacus- ochracea
azurea (mm) (mm) tris (mm) (mm)
fovea O25 0175" uv O125 DIS uva 7025-015 juve) :09925520:2
II 0.175- 0.2 I 0.175 - 0.2 Il 0.25 - 0.3
IT 02-0275 I 0252025 II 0.25 - 0.35 II 0.375 - 0.475
MV 0325-04 IV 0.375-0.475 IV 0.4 - 0.525 IV 0.625 - 0.825
V 0.5 - 0.675 V 0.575 - 0.75 V 0.675 -0.875 V OOS) Sal A275)
142
= 10%
Relative abundance of size classes (%)
RUTH BANZIGER
31.V.94 Caenis horaria, n = 68
C. luctuosa, n = 45
nine
04.V11.94
Caenis horaria, n = 3
C. luctuosa, n = 56 | | E
05.V111.94 Caenis horaria, n = 87
| C. luctuosa, n = 116
4 | IT Pes
26.V111.94
Caenis horaria, n = 335
C. luctuosa, n = 334
19.1X.94
TIR
31.X.94
i Pdl | | | Île
29.X1.94
alli nino
8 ‘OR Ne
2 = 6
Caenis horaria, n = 435
C. luctuosa, n = 693
Caenis horaria, n = 319
C. luctuosa, n = 203
Caenis horaria, n = 279
C. luctuosa, n = 207
2 14 16 18 Size classes
AQUATIC INSECTS IN MACROPHYTE BEDS 143
Combining his results and our measurement yielded the following correspondence
between HCWs and instars. Instar I: 0.2-0.3 mm; H: 0.325-0.5 mm; III: 0.525-0.775;
IV: 0.8-0.925; V: 0.95-1.1 mm. The different instars are overlapping.
Emergence time of larvae and nymphs was based on numbers and size of the
larvae and nymphs at the different sampling sessions.
RESULTS
COMPARISON OF GROWTH AND EMERGENCE TIME OF CONGENERIC SPECIES
The three pairs of congeneric species Caenis spp., Mystacides spp. and Oecetis
spp. showed staggering in emergence patterns. This delay was already prepared for in
autumn, C. luctuosa, M. longicornis and O. lacustris did stop growing from the middle
of September, whereas, C. horaria, M.azurea and O. ochracea showed some growth
until October - November.
According to the low numbers catched beginning of July, imagos of Caenis
horaria emerged between end of May and June. Juveniles appeared in the samples at
the beginning of August, but their abundance was highest at the end of August (Fig. 1).
At that time, the new generation extended over one month divided into two density
peaks: the first one (low density) ranging from size classes 9 to 15 and the second one
(high density) ranging from 2 to 8. The nymphs from the first (9-15) peak emerged by
the middle of September. Nymphs of the second peak grew until the end of October.
Most of them entered the winter period at size classes from 8 to 11.
Caenis luctuosa emerged later than C. horaria: at the beginning of July.
Juveniles were collected in the samples at the beginning of August, as for C. horaria.
The two species followed the same growth schedule (with two size class peaks at the
end of August) until the middle of September. At that time C. luctuosa stopped
growing, thus entering the winter period at size classes from 3-8 essentially.
Interpretation of the growth of Mystacides azurea was more difficult as
sampling in May seemed to have bypassed the emergence of M. azurea (Fig. 2). M. lon-
gicornis emerged at the beginning of July and the first hatched larvae of M. azurea
were ready to emerge at the beginning of August. Some M. longicornis larvae did also
reach instar V at the end of August and emerged or disappeared until the middle of
September. By the end of October, instars II to IV of M. longicornis and III and IV of
M. azurea were present. M. longicornis did not grow further, whereas M. azurea was
found from instars II to V at the end of November. Entering the winter in the last instar
could preclude to an early emergence in spring.
In Oecetis spp., emergence time extended from the end of May through August
for O. ochracea, while the emergence of O. lacustris was observed from beginning of
July through August. Some of the first hatched larvae of the two species appeared at the
Fic. |
Size classes and headcapsule widths of Caenis horaria (open columns) and C. luctuosa (dark
columns). Black arrows: emergence.
144
RUTH BÄNZIGER
beginning of August. O. lacustris did not grow further from the middle of September
and overwintered in the instars II and III, while growth of O. ochracea lasted until the
end of October and it overwintered in the instars III and IV.
GROWTH AND EMERGENCE OF ACENTRIA EPHEMERELLA
End of May was the end of the emergence period for A. ephemerella (Fig. 2).
There were still some pupae found. At the beginning of July, the first new larvae
Instars (
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Fic. 2. Frequency of the instars of five insect species (Mystacides azurea, M. longicornis,
Oecetis lacustris, O. ochracea and Acentria ephemerella).
Juv.: indetermined juveniles of
either Mystacides spp. or Oecetis spp. Black arrows: emergence.
AQUATIC INSECTS IN MACROPHYTE BEDS 145
appeared and at the beginning of August, all larval instars were present. The question
remained whether instar V larvae were the first hatched of the 1994 cohort or the last
hatched of the 1993 cohort. From August to November, young larvae grew to instar II
and III and instar IV larvae disappeared. Abundance of instar II and IH larvae was
similar at the end of November.
SPATIAL DISTRIBUTION OF INSTARS
None of the seven insect species studied showed significant differences in
relative abundance of the different instars or size classes between edge and centre of the
macrophyte bed and the adjacent substrate. Our data and figures did however show
higher relative abundances of instar V larvae on the plants (edge and centre), especially
for Oecetis spp. and Mystacides spp. (Fig. 4).
Caenis spp. showed no tendency to colonize either plants or adjacent sediments
(Fig. 3) and Acentria ephemerella was found quite exclusively on the plants (Fig. 5).
M. azurea was slightly more abundant on the adjacent sediments and on the plants in
the edge at the beginning of August, whereas M. longicornis was more abundant in the
centre at instar V. Thus, it seemed to emerge in higher densities from the centre of the
macrophyte bed than M. azurea.
Oecetis spp. were the only species which were quite abundant in the sediments.
In May, they were mostly emerging from the sediments and in the subsequent sampling
periods they were present in higher densities on the macrophytes, mainly at instar V.
The bulk of emergence of O. lacustris originated from the edge, while O. ochracea
emerged indifferently from the edge or the centre of the macrophyte bed.
Acentria ephemerella showed highest densities throughout the sampling sessions
in the centre of the macrophyte bed, but at the end of May it was mainly found in the
edge, besides of the pupae which were found in the centre. Only one individual was
found at the beginning of August on the adjacent sediments.
DISCUSSION
The distribution of larval instars and size classes of the investigated congeneric
insect species showed that temporal spacing of emergence was a major factor involved
in their segregation. Indeed, several authors pointed out the importance of separated
swarming in order to optimise mating success (Brittain, 1982). Moreover, the instar
segregation already took place in autumn which allows to minimise resource depletion
by similarly sized and similarly feeding larvae in winter when food sources are scarce.
In Oecetis spp., this difference was enhanced by the lower size of O. lacustris (usually
instar V larvae of O. lacustris had the size of instar IV larvae of O. ochracea). Size
differences between species were lower for the two other congeneric species studied
(Mystacides spp. and Caenis spp.).
The spacing of emergence involved that some species (1.e. Oecetis spp.) emer-
ged before macrophyte resumed growth and they therefore did not need the presence of
plants to complete their development. However, once the macrophytes were well
established, Oecetis spp. and Mystacides spp. were found on macrophytes prior to
RUTH BÄNZIGER
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AQUATIC INSECTS IN MACROPHYTE BEDS 147
emergence. This may be ought to the advantage of the closer distance to the air-water
interface which makes emergence easier and reduces the danger of predation (Rooke,
1984).
Among the seven species studied only two are known to mostly rely on macro-
phytes for feeding: Mystacides longicornis and Acentria ephemerella. Thus the main
food resource in relation with macrophytes (1.e. plant tissue and periphyton) was not
competed for to complete development (Berg, 1941; McGaha, 1952; Lepneva, 1966).
The absence of difference in spatial distribution of congeneric taxa may either indicate
that there is no spatial segregation or that it occurs at the microhabitat scale, as
observed by Hildrew & Edington (1979) in rivers, rather than at the edge and centre
scale of macrophyte beds. As Magdych (1979) and Miiller-Liebenau (1956) reported,
spatial segregation may take place between leaves, stems and roots at different heights
(top, middle, bottom) of the plant.
Some instar I larvae of Oecetis spp. and Mystacides spp. may have been over-
looked, thus biasing abundance data on the distribution of juveniles (Bass et al., 1982).
Caenis spp. showed higher densities on the sediments. They emerged from any
substrate and except for emergence and hibernation, their size classes were synchro-
nised. The separation of the newly hatched nymphs into a fast growing (emerging after
two months) and slow growing (hibernating) generation was already evidenced for
C. horaria by Oertli (1992) in ponds of the Geneva area. Landa (1968) observed the
same pattern for C. horaria during its study on central Europe Ephemeroptera. The
variable life cycle patterns of Caenidae were reported by many authors (see Clifford,
1982). Thus, it is interesting to note at least identical patterns in different years and
biotopes, but in the same area (i.e. Geneva), for one species.
Oecetis spp. illustrated the movement of larvae between the sediments and the
macrophytes. Last instar larvae were more abundant on the sediments in May than in
the subsequent sampling periods. It is likely that the larvae of O. ochracea did not have
enough time (or food) to colonize the macrophytes early in the season and therefore
they were still found on the adjacent sediments at the time of emergence. O. lacustris
which emerged at the beginning of July, was able to emerge partly from the macro-
phytes. O. ochracea seemed to be clearly bivoltine, whereas O. lacustris showed a
mainly univoltine pattern. These two species are at least partly predators so they are not
relying on macrophytes for feeding (Mackay & Wiggins, 1970).
M. longicornis, which was found in substantially higher densities on macro-
phytes than on sediments (Bänziger, 1998), seemed to be synchronised with macro-
phyte growth. It emerged only once the macrophytes were established, and newly
hatched larvae seemed to grow fast enabling them to emerge between the end of August
and September. Thus this species showed a fast summer generation and a slower
growing winter generation. M. azurea was more abundant on adjacent sediments than
on macrophytes. It seemed to emerge early and grow slowly, as a limited number of
larvae from the year emerged until autumn. Fast growth of M. longicornis and slow
growth of M. azurea were also reported from Petersson (1989) in southern Sweden.
This study showed that closely related taxa with similar ecological niches may
coexist in macrophyte beds at some developmental stages. Thus competition seemed
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Frequency of different larval instars of Acentria ephemerella in the edge and in the centre of
the macrophyte bed and on the adjacent sediment.
not to be involved in species distribution during most of the invertebrate life cycle.
Several papers dealing with invertebrate distribution on macrophytes in lakes conclude
at an absence of competition in this habitat (Magdych, 1979). Hargeby (1990)
explained this absence of competition by the yearly disturbance undergone by the
invertebrates through the annual life cycle of the macrophytes. Each year colonization
has to be resumed and competition has not enough time to take place (Pickett & White,
1985).
150 RUTH BÄNZIGER
Temporal segregation occurred, however, between congeneric species before
hibernation and during emergence. It may allow better resource partitioning when
resources are scarce in winter and when maximum energy and food is needed just
before emergence. Though there was a tendency for some taxa to live on the edge or on
the adjacent sediment at small size classes and to be more abundant in the centre at the
last instars or bigger size classes, this (statistically unconfirmed) result may be biased
by the impact of predation at the edge on large and moving insects.
ACKNOWLEDGEMENTS
I am grateful to all friends and colleagues who helped with the field work. In the
laboratory, I thank B. Oertli for his stimulating discussions and D. Cambin who helped
with HCW measurements. Special thanks to J.-P. Haenni who provided informations on
A. ephemerella. | am grateful to B. Lods-Crozet and E. Castella for critical review of
earlier drafts of this paper. Production of this paper was possible thanks to a grant from
the Academic Society of Geneva.
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N
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REVUE SUISSE DE ZOOLOGIE 107 (1): 153-163; mars 2000
Les Nanophthalmus Motschulsky d’Europe
(Coleoptera, Scydmaenidae)
Claude BESUCHET! & Stanislav VIT?
| Muséum d’histoire naturelle, case postale 6434, CH-1211 Genève 6.
2 Rue de la Poterie 26, CH-1202 Genève.
European species of the genus Nanopthalmus Motschulsky (Coleoptera,
Scydmaenidae). - Two new species of the genus Nanophthalmus Motschul-
sky, 1851, N. nonveilleri sp. n. et N. serbicus sp. n. are described from Yugo-
slavia (Serbia), N. ditomus Saulcy, treated previously as synonym of N.
megaloderoides Motschulsky is revalidated and the lectotype of the species
designated. Additional data related to N. megaloderoides Motschulsky,
N. turcicus Reitter and N. beszedesi Reitter are provided together with a key
to all treated species.
Key words: Coleoptera - Scydmaenidae - Cephenniini - Nanophthalmus -
taxonomy - Europe.
INTRODUCTION
Seulement sept especes ont été décrites dans le genre Nanophthalmus Mot-
schulsky, 1851: megaloderoides Motschulsky, 1851 de Crimée (espece-type); ditomus
(Saulcy, 1878) du Caucase; rotundicollis (Reitter, 1881) de Talysch; armeniacus
Reitter, 1884 du Caucase: turcicus Reitter, 1894 de Turquie d'Europe; robustus Roubal,
1913 du Caucase et beszedesi Reitter, 1913 de l’Istrie. Mais plusieurs espèces nouvelles
ont été découvertes au cours des vingt dernières années, par le Professeur Guido
Nonveiller en Serbie, par M. Rudolf Rous dans le Caucase, par M. Antoine Senglet
dans le nord de l'Iran, et enfin par nous en Turquie d’Asie. Des lacunes importantes
sont ainsi comblées dans l’aire de répartition du genre Nanophthalmus.
Nous avons commencé la révision de ce genre, devenue indispensable. Mais
l’échéance du “Catalogue of the Palearctic Coleoptera”, nous contraint à une première
mise au point, à savoir la redescription des Nanophthalmus megaloderoides Mot-
schulsky, turcicus Reitter et beszedesi Reitter, la revalidation de N. ditomus Saulcy et la
description de deux espèces nouvelles de Serbie. Ces six Nanophthalmus appartiennent
au groupe d’espèces de petite taille, ne dépassant en general pas 1 mm.
Manuscrit accepté le 09.02.1999
154 CLAUDE BESUCHET & STANISLAV VIT
MATERIEL ET METHODES
Abréviations utilisées:
HNHM Hungarian Natural History Museum, Budapest, Hongrie
IPVB Institut pour la Protection des Végétaux, Belgrade, Yougoslavie
MHNG Museum d’histoire naturelle, Geneve, Suisse
MNHN Museum National d’Histoire naturelle, Paris, France
MCSN Museo Civico di Storia Naturale “Giacomo Doria”, Gênes, Italie
MSNT Museo Civico di Storia Naturale, Trieste, Italie
NHMW Naturhistorisches Museum, Wien, Autriche
SMNS Staatliches Museum für Naturkunde, Stuttgart, Allemagne
SMFD Forschungsinstitut und Naturmuseum Senckenberg, Frankfurt/Main,
Allemagne
ZMHB Museum fiir Naturkunde der Humboldt Universitat, Berlin, Allemagne
Autres abréviations: A. - antennes; E. - élytres; Lo. - longueur; La. - largeur: P. -
pronotum; les étiquettes des matériaux étudiés sont citées ici de maniere strictement
conforme a l’original:/.../.- étiquette citée: /”....”/ - passages manuscrits de I’ étiquette.
DESCRIPTIONS
Genre Nanophthalmus Motschulsky, 1851 Biesse?
Nanophthalmus Motschulsky, 1851: 506; espèce-type Nanophthalmus megaloderoides Mot-
schulsky, 1851, par monotypie.
Cephennium (Nanophthalmus), Reitter, 1881: 554
Nanophthalmus, Cziki, 1919:17
Nanophthalmus, Lazorko, 1962: 277
Nanophthalmus, Newton & Franz, 1998: 143
DIAGNOSE. Scydmaenidae Cephenniini anophtalmes et aptères. Téte petite, les tempes
saillantes. Antennes (Fig. I) de 11 articles, la massue bien développée, formée de deux
articles seulement. Pronotum nettement convexe, un peu moins large que les élytres;
côtés arrondis, finement rebordés; aucune ornementation particulière sur le pronotum;
pas d’apophyse prosternale. Elytres convexes, sans carène laterale, ornés chacun d’une
petite fossette basale profonde, tomenteuse, accompagnée du còté externe d’une caré-
nule très courte; pas de carène humérale à proprement parler mais un pli normalement
caché sous la base du pronotum; apex des élytres souvent rebordé. Edéage formé d’une
grande capsule basale plus ou moins nettement prolongée par une sorte de manchon
dans lequel se trouve l’armure copulatrice; paramères gréles, atteignant l’apex de
l’édéage, terminés par une soie.
Nanophthalmus megaloderoides Motschulsky, 1851 Fe
Nanophthalmus megaloderoides Motschulsky, 1851: 506
Nanophthalmus megaloderoides; Motschulsky, 1869: 271;
Cephennium (Nanophthalmus) megaloderoides; Reitter, 1881: 555;
Cephennium (Nanophthalmus) megaloderoides; Reitter, 1884:84;
Nanophthalmus megaloderoides: Lazorko, 1962: 314-316, fig. 15;
LES NANOPHTHALMUS MOTSCHULSKY D'EUROPE 155
MATERIAUX EXAMINES: Type perdu. (Loc.typ. Ukraine, Aloupka). 31 ex. /laila Gebirge, Krim,
Winkler/ (HNHM, MCSN, MHNG, MSNT, NHMW, SMFD, ZMHB).
La collection Victor de Motschulsky du Muséum de Moscou ne renferme aucun
exemplaire du genre Nanophthalmus et l’exemplaire ayant servi à la description origi-
nale semble perdu. L’espece a été décrite de la Crimée (Tauride, Aloupka). Reitter
(1881) n’a pas dü voir des exemplaires de Motschulsky comme le montre sa description
qui n° est en fait qu'une mise en commun des données publiées par Motschulsky et de
Saulcy (ditomus). La diagnose ultérieure et l’édéage publiés pour l’espece megalo-
deroides Motschulsky par Lazorko (1962) s’appuient sur des exemplaires provenant des
Monts Aj Petri, pris par Moczarski et Winkler en mai 1911, qui figurent dans diverses
collections, étiquetés collectivement laila-Gebirge Krim, Winkler (ou) Moczarski. Ces
exemplaires concordent avec la description originale et appartiennent a une seule
espece, bien caractérisée par son édéage, la seule du genre Nanopththalmus connue jus-
qu'à present de la Crimée. C’est l’espèce la plus occidentale des territoires septen-
trionaux de la Mer Noire.
DIAGNOSE. Longueur (tête comprise) 0,87-0,97 mm. Elytres moins allongés que chez
les espèces balcaniques, ratio E.Lo./E.La. = 1,34-1,41; extrémités apicales des élytres
faiblement modifiées, bords épaissis sans être nettement relevés. Pronotum sur la ligne
médiane à peu près aussi long que large à la base; côtés du pronotum arqués,
convergeant en arrière vers les angles postérieurs; base droite, ratio P.Lo./P.La. = 0,78-
0,88; largeur du pronotum subégale ou inférieure à la longueur des antennes; dépres-
sions basales des élytres effacées: angles huméraux avec une fine carène prolongée en
arrière.
Antennes: ratio A.Lo./ P.base = 1,14-1,34; articles 3-8 courts, suboblongs ou
sphériques, article 9 petit et subsphérique chez les males, plus grand, pyriforme et plus
transverse chez les femelles; article 11 allonge.
Face ventrale: carénules du métasternum courtes et écartées.
Caracteres sexuels des males. Protibias recourbés et plus brusquement renflés
dans la moitié apicale; face ventrale de la portion basale du profémur pourvue d’une
fine carene terminée distalement par une minuscule saillie obtuse; métasternum ample-
ment déprimé (aplati chez la femelle).
Edéage (Fig. 3).
DISTRIBUTION. Ukraine, Crimée méridionale: Iaila Gebirge = Monts Aj Petri (nom
actuel). Aloupka = Alupka, la localité typique, se trouve sur le versant sud de cette
chaine monagneuse.
Nanophthalmus ditomus (Saulcy, 1878) Fig. 4
Cephennium ditomum Saulcy, 1878: 139-140, pl. II, fig. 21
Cephennium (Nanophthalmus) megaloderoides Motschulsky = ditomum Saulcy; Reitter, 1881:
555
Cephennium ditomum Saulcy = ? Cephennium (Nanophthalmus) megaloderoides Motschulsky;
Lazorko, 1962: 316
MATERIEL ETUDIE. Lectotype (par présente désignation) d : (sans provenance), /disque or/, /”dito-
mum n. sp. Saulcy, Typ.’/,/" 14” (étiquette bleue et noire)/, /Typus (étiquette rouge)/, MUSEUM
156 CLAUDE BESUCHET & STANISLAV VIT
DE PARIS 1900 coll. J. CROISSANDEAU/. (MHNP); 14, 12, /disque or/, /disque bleu/
” Caucase” (étiquette rose)/, /"Cephenn. ditomum Slcy - Cauc (Rtt)”/, /”14” (étiquette bleue et
noire)/, /(étiquette rouge vide)/, MUSEUM DE PARIS 1900 coll. J. CROISSANDEAU/; 16,
/disque or/, /Caucasus, Meskisches Geb. Leder, (Reitter)/, /’Cephenn. ditomum Saulcy,
Caucas”/,/"29” (étiquette bleue et noire)/, /(étiquette bleue)/, MUSEUM DE PARIS 1900 coll. J.
CROISSANDEAU/; 1d, /disque or/, /Caucasus, Meskisches Geb. Leder, (Reitter)/, /°14”
(étiquette bleue et noire)/, MUSEUM DE PARIS 1900 coll. J. CROISSANDEAU/ (tous les
exemplaires MNHN). Autres:8¢d, 1222, /Caucasus, Meskisches Geb. Leder (Reitter)/,
(MNHN, MHNG, NHMW, HNHM, MCSN).
Nanophthalmus ditomus Saulcy est une bonne espece. Nous avons pu examiner
des matériaux de l’époque, retrouvés dans la collection Croissandeau, et constater que
la mise en synonymie de cette espece par Reitter (1881:555) n’était pas fondée. Faute
de matériel disponible de Suram, localité-type de ditomus, Lazorko (1962:316) main-
tient formellement la synonymie de Reitter tout en soulignant son caractere douteux.
Nous avons choisi pour lectotype un male de la collection Croissandeau du Muséum de
Paris, portant l’étiquette manuscrite de Saulcy (voir ci-dessous).
DIAGNOSE. Longueur (téte comprise) 0,92-0,99 mm (Saulcy 0.75 mm!). Elytres moins
allongés, ratio E.Lo./E.La. = 1,35-1,45; extremites apicales des élytres modifiées, bords
faiblement rebordés et relevés. Pronotum sur la ligne médiane plus court que large a la
base; côtés du pronotum arqués, convergeant en arrière vers les angles postérieurs; base
droite, ratio P.Lo./P.La. = 0,81-0,86; largeur du pronotum généralement légèrement
supérieure ou tout au plus égale a la longueur des antennes; dépressions basales des
élytres pratiquement indistinctes; angles huméraux tout au plus avec une tres courte
carene.
Antennes distinctement plus longues que le pronotum sur la ligne médiane ou sa
largeur basale, ratio A.Lo./P.base = 1,05-1,19; articles 3,4 et 5 suboblongs, 6,7 et 8
courts, subsphériques; article 9 généralement assez gros, subpyriforme et transverse
chez le deux sexes; article 11 un peu plus long que large.
Face ventrale: carénules du métasternum courtes et écartées.
Caracteres sexuels des males. Protibias faiblement recourbés, apex avec une
protubérance dentiforme; face ventrale de la portion basale du profémur avec une fine
carene s’estompant distalement; métasternum amplement concave (faiblement convexe
chez la femelle).
Edéage (Fig. 4).
DISTRIBUTION. Géorgie, Suram (localité-type), situé à l'extrême nord-est de Meshedskiy
Khrebet (= Meskisches Gebiet des auteurs allemands).
Nanophthalmus turcicus Reitter, 1894 His
Nanophthalmus turcicus Reitter, 1894: 114.
MATERIEL ETUDIE. Holotype d étiquetté:/’Nanophthal. turcicus m. 1894”/, /’Eur. Turkei
Merkl.”/, /disque or/, MUSEUM DE PARIS 1900 coll. J. CROISSANDEAU/. (MNHN)
/Typus/”tureicus Rtt.” Cl. Besuchet det. XII. 1957/. (MHNP); 26 8, 12 / “Belgrader Wald près
Istanbul, 8.V.58, H. Schweiger”/. (MHNG).
DIAGNOSE. Longueur (tete comprise) 0,94-1,05 mm (Reitter 0,8 mm!). Elytres moins
allongés, ratio E.Lo/E.La. = 1,28-1,38; extrémités apicales des élytres a peine
LES NANOPHTHALMUS MOTSCHULSKY D’EUROPE 157
Fics 1-4
l. Nanophthalmus sp., antenne; 2. Nanophthalmus sp., apex du protibia; 3. Nanophthalmus
megaloderoides Motschulsky, édéage face ventrale; 4. Nanophthaimus ditomus (Saulcy), édéage
face ventrale. (Echelle en mm).
158 CLAUDE BESUCHET & STANISLAV VIT
modifiées, simplement renflées. Pronotum sur la ligne médiane aussi long que large à la
base; cotés du pronotum arqués, convergeant en arriere vers les angles postérieurs; base
droite, ratio P.Lo./P.La. = 07,8-0,82; largeur du pronotum légèrement inférieure à la
longueur des antennes; dépressions basales des élytres pratiquement indistinctes; angles
huméraux tout au plus avec une très courte carène.
Antennes très nettement plus longues que le pronotum sur la ligne médiane ,
ratio A.Lo./P. base = 1,13-1,28; articles 3 et 4 subégaux, oblongs, 7 et 8 courts,
subsphériques; article 9 plus gros, transverse chez le deux sexes; article 11 moins d’une
fois et demie plus long que large.
Face ventrale: carénules du métasternum courtes mais peu écartées.
Caracteres sexuels des males: protibias faiblement recourbés et renflés dans leur
moitié apicale; crochet apico-interne petit, peu distinct; face ventrale de la portion
basale du profémur pourvue d’une fine carene terminée distalement par une minuscule
saillie obtuse; métasternum amplement concave chez le male, simplement applati chez
la femelle.
Edéage (Fig. 5).
DISTRIBUTION. Turquie, Thrace, Forét de Belgrade pres d’Istanbul.
Nanophthalmus beszedesi Reitter, 1913 Fig. 6
Nanophthalmus beszedesi Reitter, 1913: 140-141.
MATERIAUX EXAMINES. Lectotype (par présente désignation) 9: étiqueté, /Dr. v. Beszédes, M.
Maggiore, Istrien, “15/8°/ Mus. HMHN det. Reitter Nanophthalmus beszedesi. Paralectotype (par
présente désignation) 1 ©, /idem/, (les deux HNHM). Il s’agit bien des matériaux vus par Reitter
qui cite “2 ex. am 15 August v. J.”. Autres: 1¢, /Dr. Beszedes, M. Maggiore, Istria “22.V.
1913”/, /’Nanophthalmus Beszedesi Reitter”/. (MHNG); 14, /idem/. (MCSN.- coll Dodero); 14,
/Mte Maggiore, Istr. Winkler. (ZMHB); 3 ex. /Mte Maggiore, Winkler/. (SMFD).
DIAGNOSE. Longueur (tête comprise) 0,87-0,92 mm. Elytres très allongés, ratio E.Lo./
E.La. = 1,48-1,53; extrémités apicales des élytres tres nettement modifiees, présentant
un fort bourrelet retroussé vers le haut, échancré a la suture. Pronotum sur la ligne
médiane aussi long que large a la base: côtés du pronotum fortement arqués au tiers
antérieur, subparallele en arriere avant les angles postérieurs; ratio P.Lo./P.La. = 0,87-
0,88; largeur du pronotum égale ou inférieure à la longueur des antennes; callosités
humérales distinctes; dépressions basales indistinctes.
Antennes nettement plus longues que le pronotum sur la ligne médiane ou
comme sa largeur à la base, ratio A.L/ P.base = 1,14-1,2 ; articles 3, 4 et 5 à peine
oblongs, 6, 7 et 8 subsphériques, article 9 petit, subsphérique chez le mâle, plus grand et
pyriforme chez la femelle: article 11 court, tout au plus d’un quart plus long que large.
Face ventrale: carénules mésosternales prolongeant la lame sternale à peine déve-
loppées.
Caractères sexuels des mâles. Protibias recourbes, brusquement renflés dès le
milieu, apex avec une protubérance dentiforme; face ventrale de la portion basale du
profémur sans caractères particuliers; métasternum amplement déprimé (aplati chez la
femelle).
Edéage (Fig. 6).
DISTRIBUTION. Croatie, Istrie, Monte UËka, (= Monte Maggiore).
LES NANOPHTHALMUS MOTSCHULSKY D'EUROPE 159
Nanophthalmus nonveilleri sp. n. Figs 78, 11
MATERIAUX EXAMINES. Holotype. d, étiqueté: /Serb. Tara pl., 1000 m, “11.06. 84”, Nonvll./
(IPVB); Paratypes. 14, 29 2, / idem “11.06.84”/. (MHNG); 28 d, 29 2,./ idem, “23.05. 84”/
(IPVB, MHNG); 16, / idem, “6.10. 78”/ (IPVB); 1d, /Serb. Tara pl., 900 m, “6.10. 78/ (IPVB).
16 /Serb. Tara pl., 900 m, “6.10.78/ (IPVB). 14, 29 2, /Yougoslavie, Serbie: Tara Planina,
Mitrovac (route Mitrovac-lac d’accumulation), 900 m, 23.V.1984; tamisage de feuilles mortes
dans une forêt de hétres, érables et épicéas (Besuchet)/. (MHNG).
DIAGNOSE. Longueur (tête comprise) 0,89-1,02 mm. Elytres très allongés, ratio Long./
Larg. = 1,43-1,55; extrémités apicales des élytres non modifiées, simples. Pronotum sur
la ligne médiane plus long que large à la base; côtés du pronotum fortement arqués au
milieu, convergeant en arrière nettement vers les angles postérieurs; base droite; ratio
P.Long./P.Larg. = 0,86-0,91; largeur du pronotum supérieure a la longueur des an-
tennes; base des élytres sans trace de dépression basale ou de callosité humérale.
Antennes un peu ou nettement plus longues que le pronotum sur la ligne
médiane et tres nettement plus longues que la largeur basale de celui-ci, ratio A.L/
P.base = 1,09-1,34; articles 3, 4 et 5 oblongs, 6, 7 et 8 subsphériques, article 9 trans-
verse, pyriforme, plus nettement transverse semble-t-il chez les males et plus allonge
chez les femelles: article 11 court, un peu plus long que large.
Face ventrale: carénules du métasternum faiblement divergentes.
Caracteres sexuels des males. Protibias recourbés, nettement renflés dans la
moitié distale, pourvus de crochet apical; face ventrale de la portion basale du profemur
avec une fine carene s’estompant distalement; metasternum peu differencie, a peine
aplani chez le male (faiblement convexe chez la femelle).
Edéage. (Figs. 7, 8).
DISTRIBUTION (Fig. 11). Serbie occidentale:Tara planina ( (8) localité-type). Selon Dr.
G. Nonveiller (communication personnelle) cette espece se rencontre aussi a Goc
planina, Beli izvori et Dobre vode (9); Ivanjıca, Mucina voda (10); Ovcar Banya,
Ovcarsko-Kablarska klisura, Debela Gora (11). Elle va de 600 a 1000 m d’altitude.
(Ces exemplaires ont été capturés et determinés par MM. Nonveiller et Pavicevic et
sont déposés a IPVB).
Nanophthalmus serbicus sp. n. Bios) 910-3
MATERIAUX EXAMINES. Holotype: d, étiqueté /Serb. Stol pl., 800 m, Nonvll., “26.05. 198477.
(IPVB). Paratypes: 16, /idem “26.05.1984”/. (MHNG); 2 ex. /idem ‘ ‘10.04.1987 1 (IPVB,
MNHG); 38 d, 22 9, /idem “15.5.1982” (IPVB, MHNG); 14 /Serb. Rtanj pl. “12.III. 1977.”/
(MHNG): 96 d,79 2, / Yougoslavie, Serbie: Stol Planina (30 km au nord de Bor), 900 m, 26.V.
1984; tamisage de feuilles mortes au pied de rochers, dans une forét de hétres, (Besuchet).
(MHNG); Tel 39 2,/ même provenance, 27.V.1984; lavage de terre au pied de vieilles souches
de hétres (Besuchet). (MHNG); 26 8, 12, /Yougoslavie, Serbie: Rtanj Planina, 600 m, 27.V.
1984: tamisage de feuilles mortes au fond d’un ravin, dans la forét de hétres (Besuchet).
(MHNG).
DIAGNOSE. Longueur (téte comprise) 0, 92-0,99 mm. Eytres nettement allongés, ratio
Long./Larg. = 1,39-1,5; extrémités apicales des élytres a peine modifiées, bords épaissis
sans €tre nettement relevés. Pronotum sur la ligne médiane plus court ou tout au plus
aussi long que large à la base; côtés du pronotum fortement arqués dans le tiers anté-
160 CLAUDE BESUCHET & STANISLAV VIT
Figs 5-10. Nanophthalmus, édéage face ventrale - 5. Nanophthalmus turcicus Reitter, (Holo-
type); 6. Nanophthalmus beszedesi Reitter; 7. Nanophthalmus nonveilleri sp.n., (Paratype), 8.
variabilité de l’apex de l’édéage (Paratype); 9. Nanophthalmus serbicus sp.n. (Paratype, Rtanj
planina), 10. variabilité de l’apex de l’édéage (Paratype, Stol planina). (Echelle en mm).
LES NANOPHTHALMUS MOTSCHULSKY D’ EUROPE 161
HONGRIE
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6
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Fic. 11
Carte de distribution - espèces de la Serbie: Nanophthalmus serbicus sp. n. (No. 1 - 7); Nano-
phthalmus nonveilleri sp. n. (No. 8 - 11).
162 CLAUDE BESUCHET & STANISLAV VIT
rieur, presque subparalleles avant les angles apicaux, base droite, ratio P.Long./ P.Larg.
= 0.81-0,86; largeur du pronotum subégale ou légèrement inférieure a la longueur des
antennes; angles huméraux oblitérés, nuls; dépressions basales des élytres faibles.
Antennes tres nettement plus longues que le pronotum sur la ligne médiane et
nettement plus longues que sa largeur basale, ratio A.L/ P.base = 1,08-1,33; articles 3, 4
et 5 oblongs, 6, 7 et 8 subsphériques, article 9 petit, subsphérique à faiblement trans-
verse chez les males, plus grand, pyriforme et plus nettement transverse chez les
femelles, article 11 court, pratiquement aussi large que long.
Face ventrale: carénules du métasternum fortement divergentes.
Caracteres sexuels des males. Protibias plus nettement recourbés dans leur
portion apicale seulement, nettement renflés des le milieu, dépourvus d’un crochet
apical; face ventrale de la portion basale du profémur avec une fine carene s’estompant
distalement; métasternum fortement déprimé, concave (applati chez la femelle,).
Edéage (Figs. 9, 10).
DISTRIBUTION (Fig 11). Serbie orientale:(1)Stol planina, (localité-type) et (2)Rtanj
planina;. Selon Dr. G. Nonveiller (communication personnelle) cette espece se ren-
contre aussi à Zajecar- Lenovac (3); Homolje - Ceremosnja (4); Mts. Homolje -
Mejdan Kucana (5); Jastrebac planina - Ravniste (6) et Kragujevac - Zezelj (7). Elle va
de 450 à 1200 m d altitude. Ces exemplaires ont été capturés et determines par MM.
Nonveiller et Pavicevic (déposés a IPVB).
TABLEAU DES ESPECES CITEES
Ce tableau n’a qu'une valeur indicative; les identifications doivent être contrôlées par
l’édéage.
| Apex des élytres orné d’un bourrelet fortement développé, relevé vers le
HAMSTER ER eae Vis wed os os E beszedesi Reitter
Apex des élytres non relevé, tout au plus avec un bourrelet léger........... 2
2 Elytres plus larges (0,46-0,49 mm), tres convexes; Thrace..... turcicus Reitter
: Elytres moins larges (0,40-0,43 mm), moins convexes .................... 3
3 Pubescence du pronotum et des élytres formée de soies couchées de
longueur moyenne; élytres plus courts (indice élytral 1,35-1,41)............ -
- Pubescence du pronotum et des élytres formée de soies semi-dressées
assez longues: élytres plus allongés (indice élytral 1,42-1,55).............. 5
4 Corps moins déprimé (haut. - 0,35 mm): en vue latérale la convexité du
pronotum est nettement séparée de celle des élytres; Géorgie . ditomus (Saulcy)
- Corps relativement déprimé (haut. ~ 0,30 mm): en vue latérale la
convexité du pronotum se prolonge sur celle des élytres; Crimée
A Rael Nope Varn SARE URI megaloderoides Motschulsky
5) Antennes légèrement plus longues, les articles 6 et 7 légèrement plus
longs que larges; métasternum du mâle avec une dépression arrondie
profonde3Serbie tr. 1 LIETA Oe ee ee ee serbicus sp. n.
- Antennes légèrement plus courtes, les articles 6 et 7 à peine ou lége-
rement plus larges que longs; métasternum du male avec une
dépression triangulaire très superficielle; Serbie............. nonveilleri Sp.n.
LES NANOPHTHALMUS MOTSCHULSKY D’ EUROPE 163
REMERCIEMENTS
Nous tenons à remercier ici toutes les institutions citées, pour nous avoir facilité
l'étude des matériaux des collections historiques, traités dans cette note, puis plus
particulièrement le Professeur Guido Nonveiller de Zemun et l’Institut pour la
Protection des Végétaux de Belgrade, Yougoslavie, pour nous avoir confié l’étude des
matériaux provenant de leurs recherches.
BIBLIOGRAPHIE
CSIKI, E. 1919. Scydmaenidae. /n:Schenkling, S. (ed.). Coleopterorum Catalogus, Pars 70. Berlin,
W. Junk, 106 pp.
LAZORKO, W. 1962. Zwei neue Cephennium-Arten (Col. Scydmaenidae) mit einer Übersicht der
ukrainischen Arten der Tribus Cephenniini. Entomologische Arbeiten aus dem Museum
G. Frey, Tutzing 13(2): 273-320.
MOTSCHULSKY, V. DE. 1851. Enumération des nouvelles espèces de Coléoptères II. Bulletin de la
Societe Imperiale des Naturalistes de Moscou 24(2): 479-511.
MOTSCHULSKY, V. DE. 1869. Enumération des nouvelles espèces de Coléoptères rapportés de ses
voyages (7e), Scydmaenides. Bulletin de la Société Impériale des Naturalistes de Moscou
42(1): 252-272.
NEWTON, A. F. & FRANZ, H. 1998. World catalogue of the genera of Scydmaenidae (Coleoptera).
Koleopterologische Rundschau 68: 137-165.
REITTER, E. 1881. Bestimmungs-Tabellen der europäischen Coleopteren. V. Paussidae, Clavi-
geridae, Pselaphidae und Scydmaenidae. Verhandlungen der Kaiserlich-Königlichen
Zoologisch-Botanischen Gesellschaft in Wien 31: 443-593, pl. 6, 7.
REITTER, E. 1884. Bestimmungs-Tabellen der europäische Coleopteren. X. Nachtrag zu den V.
Theile, enthaltend:Clavigeridae, Pselaphidae und Scydmaenidae. Verhandlungen der
Kaiserlich-Königlichen Zoologisch-Botanischen Gesellschaft in Wien 34: 59-94.
REITTER, E. 1894. Neue Pselaphiden und Scydmaeniden aus der europäischen Türkei. Wiener
Entomologische Zeitung 13: 113-115.
REITTER, E. 1913. Eine Serie neuer Scydmaeniden aus der europäischen Fauna. Entomologische
Blätter 9: 139-143.
ROUBAL J. 1913. Zwei neue paläarktische Coleopteren. Entomologische Mitteilungen, Berlin-
Dahlem 11(1): 21-22.
SAULCY, F. DE. 1878. (Pselaphidae, Scydmaenidae). /n: Schneider, O. & Leder, H. Beiträge zur
Kenntniss der kaukasischen Käferfauna. Verhandlungen des Naturforschenden Vereines
in Brünn (1877) 16: 3-258, pls. 1-4.
li Va j
et
| MGR Yes
Tate
REVUE SUISSE DE ZOOLOGIE 107 (1): 165-211; mars 2000
Über die Lauterzeugung der Welse
(Siluroidei, Ostariophysi, Teleostei) und ihren Zusammenhang
mit der Phylogenie und der Schreckreaktion *
Andreas HEYD & Wolfgang PFEIFFER
Zoologisches Institut, Universität Tübingen,
Auf der Morgenstelle 28,
D-72076 Tübingen, Germany.
Sound production in catfish (Siluroidei, Ostariophysi, Teleostei) and its
relationship to phylogeny and fright reaction. - Sound production by
pectoral stridulation, and sound production by swim-bladder mechanisms
were studied in 19 species of catfish from 8 families. Stridulation sounds
were recorded in air by means of a microphone and a tape recorder while
holding one of the pectoral spines; swim-bladder sounds were recorded
under water. All sounds were investigated by sonographic analysis of the
records. In addition, three species (Arius seemani, Pimelodella gracilis,
Pimelodus pictus) were studied anatomically in order to determine whether
or not they have a swim-bladder mechanism.
Sound production by stridulation was observed when the enlarged pectoral
fin spines were moved in both directions (spread and attracted) in Mystus
gulio, M. vittatus (Bagridae), Synodontis ocellifer, S. schoutedeni (Mo-
chokidae), Agamyxis flavopictus (Doradidae), and Dysichthys coracoideus
(Aspredinidae). In contrast, the following species produced sounds only
when they spread their pectorals: Arius seemani (Ariidae), Pimelodella (=
Brachyrhamdia) meesi (Pimelodidae), Dianema urostriata (Callichthy-
idae), Peckoltia pulcher, and Glyptoperichthys gibbiceps (Loricariidae). In
the other Loricariidae studied sound production was absent.
In addition to the pectoral stridulation mechanism, Arius seemani possesses
a modified swim-bladder mechanism, called the "Springfederapparat“
(Müller, 1842) or „elastic spring apparatus“. Thin elastic bones, derived
from the transverse processes of the first few vertebrae, function in sound
production. Specialized sonic muscles on the upper surface of this „elastic
spring“ cause the vibration of the swim-bladder. The elastic bones function
as antagonists of the muscular contraction. They return the fibres of the
sonic muscles after each contraction. The sonic activity of A. seemani with
its swim-bladder mechanism showed a daily rhythm. The choruses were
most vigorous in the early morning. The sounds emitted consisted of short
* In memoriam Prof. Dr. Karl von Frisch
Manuskript angenommen am 9.11.1999
166
ANDREAS HEYD & WOLFGANG PFEIFFER
pulses at irregular intervals with a fundamental frequency near 170 + 30
Hz. This frequency was equivalent to the vibration frequency of the sonic
muscles.
In Pimelodus pictus (Pimelodidae) the protractor muscle (functioning as a
vibration generator) is directly connected to the swim-bladder (serving as a
resonance body). This system generates drumming sounds. In Pimelodella
gracilis a swim-bladder mechanism could not be found.
We detected a new method of sound production in Hemibagrus nemurus
(Bagridae). It is able to squeak or to scream by pressing out air from its gill
slits.
The frequent occurence of a stridulatory mechanism by means of the
pectoral spines in at least 13 out of 33 families of catfish indicates that
those species which lack a stridulatory mechanism may have subsequently
lost it during evolution. In contrast, swim-bladder mechanisms are
supposed to have developed independently several times, since they are
differently structured. The existence of two very different sonic mecha-
nisms, a stridulatory as well as a swim-bladder mechanism, in the same
species (for example Arius seemani and Pimelodus pictus) indicates that
the sounds may have a different biological significance. The frequencies of
the nonharmonic sounds by stridulation (described as chirps, clicks, croaks,
grunts, knocks, scrapes, scratches, squeaks, rasps, thumps and so on) range
from 100 - 8000 Hz, mostly 1000 - 4000 Hz. Their main acoustic energy is
found at frequencies above the hearing range of most fish, except the
Ostariophysi. In addition, all predaceous fishing mammals and birds are
able to hear these frequencies. Thus, these sounds may serve as an impor-
tant means of warning and defence, especially since they are mostly
produced when the catfish is captured and pulled out of the water. The
noise may function in the same way as that produced by threatened rattle
snakes. In a similar manner to these poisonous snakes, the stridulating
catfish are defensive. The pectoral spines and the dorsal spine are erected
simultaneously in order to protect the fish. Since pectoral and dorsal spines
have locking mechanisms, predators can swallow these catfish only with
great difficulty. Whereas stridulatory mechanisms exist in many solitary
species, swim-bladder mechanisms are numerous in social catfish and are
supposed to serve intraspecific communication.
In all species of the mostly solitary and night active catfish that have been
studied with respect to their sound production, the fright reaction elicited
by the alarm substance from the epidermis of conspecifics is either little
developed (Mochokidae, Pimelodidae) or absent (Aspredinidae, Lori-
cariidae). On the other hand, no stridulatory mechanism has been detected
in some pelagic, schooling species of catfish possessing a distinct, well-
developed fright reaction and being active in daylight (Eutropiellus van-
deweyeri - Schilbeidae, Kryptopterus bicirrhis - Siluridae). However, since
only a few species of catfish have been studied, conclusions concerning a
connection between the existence of the chemically elicited fright reaction
LAUTERZEUGUNG DER WELSE (SILUROIDEI) 167
on the one hand and sound production by sonic mechanisms on the other
should only be drawn with caution.
Key-words: fish - catfish - sound production - stridulation - swim-bladder
mechanisms (Trommelmechanismen) - „elastic spring apparatus“
(Springfederapparat) - fright reaction (Schreckreaktion) - alarm substance
(Schreckstoff).
EINLEITUNG
Die Welse (Siluroidei) umfassen mit etwa 2500 rezenten Species in 33 Fami-
lien (Teugels, 1996) mehr als ein Drittel aller Ostariophysen (Sagemehl, 1885) und
ungefähr 10% der Teleostei (Eigenmann & Eigenmann, 1890; Regan, 1911; Berg,
1958; Greenwood er al., 1966; Roberts, 1973; Gosline, 1975; Novacek & Marshall,
1976; Briggs, 1979; Fink & Fink, 1981, 1996; Lauder & Liem, 1983; Lundberg &
McDade, 1986: Nelson, 1994). Mehr als 90% der Welsarten sind solitäre, in
Bodennähe lebende, nachtaktive, räuberische Süßwasserfische. Viele Callichthyidae,
Pangasiidae, Schilbeidae und manche Siluridae sind tagaktiv, schwarmbildend und
Kleintier- oder Pflanzenfresser. Die Loricariidae ernähren sich vorwiegend von
Pflanzen und Detritus. Einige Ariidae, Aspredinidae und Plotosidae leben im Meer
oder Brackwasser (Franke, 1985). Die Plotosidae zeigen ım Juvenilstadium ein
besonders stark ausgeprägtes Schwarmverhalten (Burgess, 1989). Als Ostariophysi
sensu stricto (Otophysi sensu Rosen & Greenwood, 1970) haben alle Welse einen
Weberschen Apparat (Weber, 1820; Wright, 1884; Bridge & Haddon, 1889, 1893,
1894; Sörensen, 1895; Chranilov, 1929; Alexander, 1964, 1965; Chardon, 1968). Er
verbindet die Schwimmblase mit dem Innenohr und dient der Steigerung der
Hörschärfe (von Frisch & Stetter, 1932). In vielen Welsfamilien gibt es Species, die
in ihrer Epidermis Schreckstoffzellen führen (Pfeiffer, 1960, 1970, 1977). Beim
Schreckstoff (von Frisch, 1938), der diesen Zellen entstammt (Pfeiffer, 1960; Smith,
1973), handelt es sich um ein Pheromon (Karlson & Lüscher, 1959), das bei Frei-
setzung durch Hautverletzung eine gesteigerte Aufmerksamkeit und eine Flucht-
reaktion der Artgenossen auslöst. Dieses Verhalten wurde von Karl von Frisch (1938,
1941a, b) an der Elritze (Phoxinus phoxinus, Cyprinidae) entdeckt und als Schreck-
reaktion bezeichnet. Während die Schreckreaktion ein Merkmal aller Ostariophysi
sensu lato, d.h. Gonorynchiformes (= Anotophysi, Rosen & Greenwood, 1970) plus
Otophysi, ist (Pfeiffer, 1967), tritt der Webersche Apparat nur bei den Otophysi auf
und ist somit das phylogenetisch jüngere Merkmal. Bei einigen Species oder Familien
mit besonderer Lebensweise wurde die Fähigkeit zur Schreckreaktion sekundär
teilweise oder vollständig rückgebildet (Pfeiffer, 1963a). So fehlt den Serrasalminae
und Mylinae nur die Fähigkeit zur Schreckreaktion (im ZNS), obwohl sie in ihrer
Epidermis Schreckstoffzellen (histologisch sichtbar) und damit Schreckstoff (im
Verhaltensexperiment nachgewiesen) führen (Pfeiffer, 1962a, 1963a; Markl, 1968).
Die Gymnotoidei und die Welsfamilien Aspredinidae und Loricariidae haben dagegen
im Laufe ihrer Stammesgeschichte auch die Schreckstoffzellen und damit den
Schreckstoff rückgebildet (Pfeiffer 1963a). Wie die meisten Knochenfische (Tele-
168 ANDREAS HEYD & WOLFGANG PFEIFFER
ostei) besitzen fast alle Welse (mit Ausnahme der Aspredinidae) Mauthner-Axone in
ihrem ZNS und damit eine Mauthner-Reaktion, mit der auch ihre Schreckreaktion
beginnt (Pfeiffer et al., 1986; Göhner & Pfeiffer, 1996).
Etwas Besonderes innerhalb der Ostariophysi ist die Fähigkeit zahlreicher
Welsarten, auf unterschiedliche Weise Laute zu erzeugen. Bei mindestens 8 Familien
kommen Trommelmechanismen vor, und noch häufiger treten Stridulationsapparate
auf. Innerhalb mehrerer Familien gibt es sogar beide Mechanismen nebeneinander.
Ein Trommelmuskelapparat oder Springfederapparat ist bekannt von den Ariidae,
Auchenipteridae, Bagridae, Doradidae, Malapteruridae, Mochokidae, Pangasiidae und
Pimelodidae (Müller, 1842, 1857; Sörensen, 1884, 1895; Tavolga, 1962; Abu-Gideiri
& Nasr, 1973; Kastberger, 1977, 1978; Kratochvil er al., 1980; Schachner & Schaller,
1982; Ladich & Fine, 1994). Die Schwimmblase verstärkt als Resonanzraum die
Vibration der mit ıhr direkt oder indirekt verbundenen Trommelmuskeln.
Bei der Stridulation werden durch Gegeneinanderreiben von Knochenele-
menten knarrende Geräusche erzeugt. Am weitesten verbreitet ist die Stridulation mit
den Brustflossen. Sie ist nachgewiesen für die Arıidae, Aspredinidae, Auchenipter-
idae, Bagridae, Callichthyidae, Doradidae, Heteropneustidae, Ictaluridae, Loricari-
idae, Mochokidae, Pangasiidae und Pimelodidae. Die Laute entstehen durch Be-
wegung des massiven Kopfes des ersten Brustflossenstrahls (Stachel), der einen
Knochenkamm mit Querrillen und Höckern trägt und in eine gebogene Kerbe
(Gelenkpfanne) im Cleithrum (Teil des Schultergürtels) eingepaßt ist. Wird dieser
Knochenkamm gegen den Boden der Gelenkpfanne gepreßt und gleichzeitig bewegt,
entstehen knarrende Geräusche (Geoffroy Saint-Hilaire, 1829; Dufossé, 1874;
Haddon, 1881; Sörensen, 1884; Villwock, 1960; Winn, 1964; Pfeiffer & Eisenberg,
1965; Agrawal & Sharma, 1965; Gainer, 1967; Abu-Gideiri & Nasr, 1973; Brousseau,
1976; Kastberger, 1977, 1978; Kratochvil et al., 1980; Kratochvil & Völlenkle, 1981;
Schachner & Schaller, 1982: Fine er al., 1996, 1997: Kaatz, 1999). Als Besonderheit
ist von einigen Sisoridae die dorsale Stridulation (wie von Geoffroy Saint-Hilaire,
1829 an Synodontis schall irrtümlich vermutet) bekannt. Hierbei reibt eine feilen-
förmige Struktur an der Rückenflossenbasis durch Vor- und Rückwärtsbewegung
gegen die gerippte Oberfläche der verwachsenen Neuratfortsätze des Wirbelkom-
plexes (Haddon, 1881; Mahajan, 1963; Alexander, 1965).
Bei allen Fischen mit besonderen lauterzeugenden Organen handelt es sich um
Knochenfische (Osteichthyes), meist Teleostei. Bereits Aristoteles erwähnt sechs
lauterzeugende Fischarten (Aristoteles, auch zitiert bei Cuvier & Valenciennes, 1840,
Müller, 1857 und Sörensen, 1884, 1895). Es handelt es sich dabei um Species aus
dem Mittelmeer. Geoffroy Saint-Hilaire (1829) beschrieb als erster die Lauterzeugung
durch Stridulation eines Welses, nämlich Synodontis schall (Bloch & Schneider,
1801) aus dem Nil (Daget er al., 1986). Seine richtige Aussage wurde von Cuvier &
Valenciennes (1840) ohne eigene Beobachtung abgestritten, doch von Johannes
Müller (1857) bestätigt. Erst knapp 100 Jahre nach Geoffroy Saint- Hilaire (1829) hat
Karl von Frisch (1923) mit der von ihm in die Sinnesphysiologie und Verhaltens-
forschung eingeführten genialen Dressurmethode das Hörvermögen für Fische nach-
gewiesen, und zwar am nordamerikanischen Zwergwels Ameiurus nebulosus, "der
LAUTERZEUGUNG DER WELSE (SILUROIDEI) 169
kommt, wenn man ihm pfeift" (wofür der geblendete Fisch mit Futter belohnt wurde).
Im Gegensatz zu den experimentellen Untersuchungen über Hörvermögen und
Lauterzeugung gehen die ersten anatomischen Befunde und Beobachtungen an den
lauterzeugenden Organen von Welsen weit mehr als 100 Jahre zurück (Müller, 1842;
Agassiz, 1852; Reissner, 1859; Dufossé, 1874; Haddon, 1881; Wright, 1884;
Sorensen, 1884, 1895, 1898; Bridge & Haddon, 1889, 1893, 1894).
Eine Tierart bedarf, um zu überleben, des Schutzes ihrer Individuen vor dem
Gefressenwerden durch Beutegreifer. Ein Schutz vor räuberischen Überfällen kann
gegeben sein durch die Warnung der Artgenossen, beispielsweise durch ein Alarm-
pheromon wie den Schreckstoff, und ist besonders bei wehrlosen, tagaktiven und
schwarmbildenden Fischen zu erwarten. Im Gegensatz zu den meisten Species der
Cyprinidae und Characiformes ist es gewöhnlich schwierig, die Schreckreaktion für
die Siluroidei nachzuweisen (Pfeiffer, 1960, 1963a, 1967). Schutz (1956) erzielte an
Ameiurus nebulosus acht und an Corydoras paleatus zwei positive Ergebnisse, ohne
sie als Schreckreaktion zu deuten, trotz seines Hinweises, daß diese Welse als Jung-
fische in Gruppen leben. Pfeiffer (1960) bestätigte die von Schutz (1956) an Ameiurus
und Corydoras erhobenen positiven Befunde und wies die Schreckreaktion der Welse
erstmalig an dem schwarmbildenden, tagaktiven Glaswels Kryptopterus bicirrhis
(Siluridae) nach. Seitdem wurde die Schreckreaktion dieser und weiterer Species
(Pangasius sutchi - Pangasiidae, Eutropiellus vandeweyeri - Schilbeidae und dem
marinen Plotosus lineatus - Plotosidae) videotechnisch analysiert (Pfeiffer er al. 1986,
Heyd & Pfeiffer, in Vorbereitung). Positive Ergebnisse liegen ferner für Angehörige
der Mochokidae und Pimelodidae vor (Pfeiffer, 1963a).
Eine völlig andere Möglichkeit des Schutzes ist die Abschreckung des An-
greifers, beispielsweise akustisch durch Stridulationslaute oder Knarren. Sie ist
hauptsächlich bei solitären, wehrhaften Species zu erwarten, wie den stacheltra-
genden, meist nachtaktiven Welsen. Es war daher naheliegend zu prüfen, ob bei
Welsen ein Zusammenhang zwischen Lauterzeugung und Schreckreaktion besteht,
wie von Pfeiffer & Eisenberg (1965) vermutet. Trotz zahlreicher Sammelreferate über
die Schreckreaktion einerseits (Pfeiffer, 1962b, 1963b, 1966, 1974, 1977, 1982:
Smith, 1977, 1982, 1986, 1992; Solomon, 1977; Liley, 1982; Stabell, 1996) und die
Lauterzeugung andererseits (Harden-Jones & Marshall, 1953; Tavolga, 1960, 1971a,
1977; Schneider, 1961, 1967; Winn, 1964; Popper & Fay, 1973, 1993; Fine et al.,
1977; Hawkins & Myrberg, 1983; Bass, 1989; Schellart & Wubbels, 1998; Tyack,
1998) wurde dieser Frage bisher keine Aufmerksamkeit geschenkt. Da sich die
bisherigen Untersuchungen auf nur wenige Arten und Familien beschränken, haben
wir weitere Species bezüglich ihrer Fähigkeit zur Lauterzeugung und deren Mecha-
nismen studiert.
MATERIAL UND METHODEN
Bezüglich ihrer Lauterzeugung wurden 19 Species aus 8 Familien der Welse
untersucht (Tabelle 1). Die Fische wurden als Jungtiere vom Großhandel bezogen;
alle waren Wildfänge, ausgenommen Corydoras paleatus. Sie wurden in 200L-
170 ANDREAS HEYD & WOLFGANG PFEIFFER
Aquarien in Leitungswasser bei 24 - 26 °C Wassertemperatur gehältert (LD 12:12).
Die Fütterung erfolgte je nach Species 1-2 mal täglich mit Flockenfutter, Futter-
tabletten, roten Mückenlarven (Frostfutter) und Rinderherz.
LAUTERZEUGUNG DURCH STRIDULATION:
Die Versuche begannen nach 6 Wochen Quarantänezeit und fanden in einem
ruhigen Raum statt. Die Welse wurden nach einer Eingewöhnungszeit von etwa einer
Stunde aus einem kleinen Transportaquarium gefischt, an einer Brustflosse und am
Körper gefaßt und in 5-10 cm Abstand vor ein Mikrophon (Uher M 517) gehalten.
Die bei Bewegung der freien Brustflosse erzeugten Laute wurden mit einem
Tonbandgerät (Uher report stereo) bei 19 cm/sec Bandgeschwindigkeit aufgezeichnet.
Die einzelnen Fische wurden am selben Tag höchstens zweimal aus dem Aquarium
genommen, wobei eine mindestens halbstündige Pause eingehalten wurde. Wenn ein
Wels keine Laute erzeugte, wurde er später zu unterschiedlichen Tageszeiten, auch
am Abend, wieder geprüft.
Mittels eines Sonagraphen wurden von den Tonbandaufnahmen Sonagramme
erstellt (Sonagraph Typ 6061, Papier Type B/65 Sonagram, Kay Elemetrics Corp.,
Pine Brook, NJ, USA), wobei die Registrierdauer 1,2 Sekunden im Frequenzbereich
160-16000 Hz betrug. Durch Darstellung der Intensitätsverteilung konnte eine
Frequenzanalyse der Laute durchgeführt werden. Die Tonbandaufnahmen wurden
möglichst vollständig in Sonagramme übersetzt, aus denen die Längen der Laute und
der Intervalle (Pausen zwischen den Lauten) bestimmt und in Millisekunden (ms)
umgerechnet wurden. Die Mebwerte für jedes Individuum und für die Summe aller
Angehörigen derselben Species wurden statistisch bearbeitet. Dabei wurde innerhalb
derselben Art nicht zwischen Ab- und Adduktionslauten unterschieden. Ferner konnte
durch Abspielen der Tonbänder mit reduzierter Geschwindigkeit die Anzahl der
registrierten Laute bestimmt werden. Der Quotient aus Lautanzahl und zugehöriger
Aufnahmedauer ergibt ein relatives Maß für die Aktivität (den Erregungsgrad) des
Versuchsfisches.
LAUTERZEUGUNG DURCH SCHWIMMBLASENMECHANISMEN:
Zur Lautaufnahme wurden die folgenden Geräte eingesetzt: Panasonic NV-FS
200 (SVHS-Videorecorder mit Long-Play-Aufnahmemôglichkeit); Videocassetten:
Sony VHS E-240; Dyn. Mikrophon Vivanco- DM 22 (Frequenzbereich 60-12000 Hz,
Impedanz 600 Ohm, Nierencharakteristik). Für die Unterwasseraufnahmen war dem
Mikrophon ein Gefrierbeutel übergestülpt und wasserdicht abgeklebt. Dadurch wur-
den die eingegebenen Frequenzen weder an Luft noch unter Wasser beeinflußt, wie
der folgende Versuch gezeigt hat: mit einem Frequenzgeber (SINE/SQUAR Oscil-
lator Li Interlab SQ 10 mit einem 50 Ohm/0,2 Watt Lautsprecher) wurden definierte
Töne erzeugt und über das Mikrophon unverpackt oder verpackt registriert. Diese
Aufnahmen wurden mit einem Frequenzzähler bzw. Frequenzanalyse-PC-Programm
(SONA-PC, B. Waldmann, Zoologisches Institut der Universität Tübingen) analy-
siert. Die Frequenz war stabil, das Mikrophon zeigte eine Richtcharakteristik. Mit
LAUTERZEUGUNG DER WELSE (SILUROIDEI) 171
zunehmender Entfernung vom Mikrophonkopf sank die Intensität der Töne, abhängig
von der jeweiligen Frequenz. Die Versuche wurden unter den folgenden Bedingungen
durchgeführt: Aquariengröße 80x50x50 cm, Wassertemperatur 27-28°C, Lichtphase
7-19 Uhr (Leuchtstoffröhre); Fütterung Ix täglich am Vormittag. Die Fische wurden
jeweils eine Woche eingewöhnt.
Versuch A: Zur Bestimmung der Tag-Nacht-Lautaktivität wurde im Abstand
von drei Wochen an zwei verschiedenen Gruppen aus je 4 Exemplaren von Arius
seemani (Gesamtlänge 205-240 mm) die Lautaktivität drei Tage (72 Stunden) lang
ohne Unterbrechung auf Videoband registriert. Der Aquarienraum wurde nur zum
Programmieren des Recorders, Wechseln der Cassetten und zur Fiitterung betreten,
um die Welse so selten wie möglich zu beunruhigen. Zur Bestimmung der Laut-
aktivität wurden die Bandaufnahmen abgehört, wobei von jeder Stunde drei 5-
Minuten-Abschnitte ausgewählt und jeweils die Laute gezählt wurden. Aus diesen an
den drei Aufnahmetagen immer zur selben Stunde ermittelten Werten (n = 9) wurde
der Median berechnet (= mittlere Lautaktivität in 5 Minuten). Dieser wurde mit 12
multipliziert und dient als geschätztes Maß für die Lautaktivität während der
betreffenden Stunde.
Versuch B: Nach Abschluß der beiden Langzeitversuche wurden nachmittags
drei Welse aus der zweiten Gruppe entnommen und die Laute des im Aquarium
verbliebenen Individuums sofort nach dieser Störung bzw. nach einer Beruhigungs-
phase von 15 Minuten registriert. Von diesen Aufnahmen wurden jeweils 20 Laute
mit dem Sona-PC-Programm in bezug auf Frequenzanteile (Fourier-Analyse) sowie
Laut- und Intervalldauer ausgewertet.
Für die vergleichend-morphologischen Untersuchungen dienten je 3 Arius
seemani (215-260 mm Gesamtlänge), Pimelodus pictus (105-110 mm) und Pimelo-
della gracilis (110-125 mm). Von jeder Species wurden nach Tötung mit MS 222
(Sigma Chemical Co.) ein Exemplar frisch präpariert und zwei in Formalin (1:10
verdünnt) fixiert. Alle Fische wurden ventral geöffnet und geprüft auf den Besitz
eines Fensters in der Muskulatur jeder Körperseite (“the lateral cutaneous areas”:
Bridge & Haddon, 1893; Alexander, 1964, 1965, 1966). Ferner wurden Lage, Größe
und Form ihrer Schwimmblase notiert. Schließlich wurde nach einem Trommel-
muskelapparat gesucht. Während der Präparation wurde mit einer ZEISS-Auf-
setzkamera M 35 am ZEISS-Standardbinokular fotographiert (Filmmaterial: Agfa
HDC 100/21, Fujichrome 50/18, Kodak 100/21).
ERGEBNISSE
STRIDULATION
Alle Species reagierten auf Berührung mit Zappeln, Schwanzschlagen, Auf-
richten der Rückenflosse sowie mit Abspreizen und Arretieren oder Bewegungen der
Brustflossen. Bei letzterem erzeugten 13 der 18 untersuchten Species Stridulations-
laute (Tab. 1, 2). Am lautesten waren die beiden Synodontis-Species, Agamyxis und
Glyptoperichthys, während Peckoltia besonders leise stridulierte. Die Angehörigen
derselben Art waren unterschiedlich erregt: die einen begannen schon beim Heraus-
fischen zu stridulieren, die anderen erst beim Anfassen, einige blieben auch bei
172 ANDREAS HEYD & WOLFGANG PFEIFFER
TABELLE |
Familienzugehörigkeit der 19 untersuchten Welsarten, in Tab. 2 und Abb. 6-9 verwendete
Abkürzungen (Abk.), Anzahl der auf Stridulation geprüften Exemplare (N) und deren Gesamt-
länge.
* es könnte sich um Peckoltia vittata (Steindachner, 1882) handeln.
** Syn. Pimelodella meesi. *** nur auf Trommelmechanismen untersucht.
Familie / Species Abk. N Länge (mm)
Ariidae
Arius seemani Günther, 1864 As 12 60-85
Aspredinidae
Dysichthys coracoideus Cope, 1874 De 5 80-100
Bagridae
Hemibagrus nemurus (Valenciennes, 1839) 4 110-122
Mystus gulio (Hamilton, 1822) Mg 4 63-67
Mystus vittatus (Bloch, 1794) Mv 3 55-62
Callichthyidae
Corydoras paleatus (Jenyns, 1842) 13 40-50
Dianema urostriata De Miranda-Ribeiro, 1912 Du 5 40-55
Doradidae
Agamyxis flavopictus (Steindachner, 1908) Af 3 53-65
Loricariidae
Ancistrus cf. dolichopterus Kner, 1854 3 55-70
Glyptoperichthys gibbiceps (Kner, 1854) Gg 3 100-120
Panaque nigrolineatus (Peters, 1877) 2 55-60
Parancistrus aurantiacus (Castelnau, 1855) | 80
Peckoltia pulcher (Steindachner, 1915) * Pp 3 73-90
Pseudacanthicus spinosus (Castelnau, 1855) 3 70-95
Mochokidae
Synodontis ocellifer Boulenger, 1900 So 3 105-130
Synodontis schoutedeni David, 1936 Ss 3 80-100
Pimelodidae
Brachyrhamdia meesi Sands & Black, 1985** Bm | 70
Pimelodella gracilis *** (Cuvier &
Valenciennes, 1840) = =
Pimelodus pictus Steindachner, 1876 4 75-90
wiederholtem Fangen ruhig. Eine hohe Stridulationsaktivität zeigten die beiden
Mystus- und Synodontis-Arten sowie Agamyxis flavopictus und Peckoltia pulcher
(Tab. 2). Der Erregungsgrad aller untersuchten Individuen lag im Durchschnitt
zwischen 0,9 und 2,5 Lauten pro Sekunde. Nur bei 5 Species stridulierten alle
Individuen; von 66 stridulierfähigen Individuen aller Species blieben 24 (36%) auch
bei wiederholten Störungen stumm (Tab. 2). Stridulation im Wasser trat nur selten
auf. Einmal lärmte ein Agamyxis im Aquarium, als er mit zwei Artgenossen in die
Quere kam. Ein Pimelodus stridulierte im Becherglas, mit dem er aus dem Aquarium
genommen worden war. Dysichthys beantwortete das Anfassen im Wasser mit
Körpervibrationen.
LAUTERZEUGUNG DER WELSE (SILUROIDEI) 173
TABELLE 2
Ergebnisse der Untersuchungen zur Stridulation (Abkürzungen der Speciesnamen vgl. Tabelle
1). Nur stridulierende Species sind hier aufgelistet.
S: Stridulationsweise (Ab = Abduktion, Ad = Adduktion).
n (N): Anzahl n der stridulierenden Individuen von allen geprüften Exemplaren (N) einer
Species; E: Erregungsgrad (Anzahl Laute / sec., Mittelwerte + Standardabweichung); L: Laut-
dauer (Medianwerte) in ms; P: Intervalldauer (Medianwerte) in ms; F: Frequenzschwerpunkte
der Laute in KHz.
L P F
Sea SÌ n(N) E
As Ab 6 (12) IR 0.5 30.2 77.4 1-3; 4-5; 10-11
Bm Ab 170) 0.9 52.8 TS 2-4; 6-8
Du Ab SO) 1.0+0.6 41.5 Wal 0.5-2; 3-4
Gg Ab 313) M 087] SS 2302 1.5-2.5; 4-5
Pp Ab 16) 1.6 + 0.6 67.9 130.2 4-6; 9-12
Af Ab+Ad 3:6) 1.4 +0.4 101.9 56.6 2-3; 5-6
De Ab+Ad 4 (5) 19122086 83.95 41.5 0.5-2
Mg Ab+Ad 3 (4) 14+1.2 45.3 34 3-4; 7-8
Mv Ab+Ad 348) 1.4 +0.4 34.9 34 3-5; 8-10
So Ab+Ad 28) Ses 10 61.35 Sell 1-2; 3.5-4.5
Ss Ab+Ad 2 (3) 29-13 49.1 23893 0.5-2
Die Stridulationslaute entstanden je nach Species auf unterschiedliche Weise.
Während Arius, Brachyrhamdia, Dianema, Glyptoperichthys und Peckoltia nur bei
Abduktion (Abspreizen) der Brustflosse Laute erzeugten, geschah dies bei Agamyxis,
Dysichthys, den Mystus- und den Synodontis-Arten bei Abduktion und Adduktion
(Heranziehen) der Brustflosse (Tab. 2). Lauterzeugung nur bei Adduktion wurde nie
beobachtet. Die Geräusche bestehen aus Serien einzelner Laute, welche wiederum aus
Einzelknacks aufgebaut sind, die bei kürzeren Lauten dichter aufeinander folgen als
bei längeren. Diese Einzelknacks sind bei Arten mit Abduktionslauten ziemlich
gleichmäßig angeordnet und die Laute einer Serie ähnlich gestaltet (Abb. 1, 2). Teil-
weise können anhand der Sonagramme die Abduktionslaute von den Adduktions-
lauten an ihrer unterschiedlichen Dauer bzw. der Dichte der Einzelknacks unter-
schieden werden (Abb. 3, 4, 5). Die Laute zeigen keinen harmonischen Aufbau,
sondern eine breite Frequenzverteilung. Die Bereiche größter Energie sind in den
Sonagrammen an den stark geschwärzten Stellen (Banden) erkennbar (Abb. 1-5). Oft
treten zwei oder drei Banden auf, wobei die niedrigste Frequenz den Hauptbereich
darstellt. Die Hauptfrequenzanteile liegen bei 500 - 6000 Hz (Tab. 2).
Aufgrund individueller Unterschiede (Abb. 6) ergeben sich in der Zusammen-
fassung für jede Species teilweise große Schwankungen der Laut- und noch mehr der
Pausenlängen (Abb. 7). Kurze Laute im Bereich bis 50 ms erzeugen Arius, Dianema,
Mystus gulio, M. vittatus sowie Synodontis schoutedeni. Bei den restlichen Species
betragen die mittleren Lautlängen 50 - 100 ms (Tab. 2). Die größte Variabilität der
Lautlänge zeigen Agamyxis, Dysichthys, Glyptoperichthys und Peckoltia, wobei ein-
zelne Laute mehr als 150 ms dauern. Auch das einzige Exemplar von Brachyrhamdia
174
ANDREAS HEYD & WOLFGANG PFEIFFER
Mm
2 4
az Be
= 2
È è iu À
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sie va ve
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ur
|
| [ll |
STILI
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È > ji LA Mi dl) A
2 2. I Ht 1 Mi
100 ms
ie
1 Be
ABB. |
Ausschnitt je eines Sonagramms von: a) Arius seemani, b) Brachyrhamdia meesi, c) Dianema
urostriata. Abszisse: Zeitskala in ms. Ordinate: Frequenzskala 0-16 kHz in 1-kHz-Stufen.
LAUTERZEUGUNG DER WELSE (SILUROIDEI) 175
| N
| |
ti);
|
ABB. 2
Ausschnitt je eines Sonagramms von: a) Glyptoperichthys gibbiceps, b) Peckoltia pulcher.
Achsen wie in Abb. 1.
weist bei kürzerer Lautdauer eine große Streuung auf (Abb. 7a). Die Pausenlänge ist
am geringsten und streut am wenigsten bei den zwei Synodontis-Arten und Mystus
vittatus; sie ist am größten bei Glyptoperichthys und streut am breitesten bei Dianema
und Dysichthys (Abb. 7b).
Während ein Zusammenhang zwischen Erregungsgrad und Laut- bzw.
Pausenlängen nicht ersichtlich ist (Tab. 2), zeigt sich im Verhältnis von Lautlänge zu
Pausenlänge eine Abhängigkeit von der Art der Lauterzeugung (Abb. 8). Bei Species,
die Ab- und Adduktionslaute erzeugen, sind die Laute mindestens ebenso lang wie die
Pausen (Verhältnis meist größer als 1); bei Species mit Abduktionslauten übersteigt
die Intervalldauer die Lautdauer teilweise weit (Verhältnis kleiner als 1).
176
ANDREAS HEYD & WOLFGANG PFEIFFER
my
ni
N |
[LI
ME
wi
m
%
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ie
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ds
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ee
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HR ©;
100 ms
ABB. 3
Ausschnitt je eines Sonagramms von: a) Mystus gulio, b) Mystus vittatus. Achsen wie in Abb. 1.
LAUTERZEUGUNG DER WELSE (SILUROIDEI) 11/7
Hi
N Il
& È DT | ev h
trad à
L'ART 4 vr. ry
n SR = dle, m nm oe
L- = a a Me
= = = eee == = =
a Meet Pr. Zi ee SII
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kr
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100 ms
ABB. 4
Ausschnitt je eines Sonagramms von: a) Synodontis ocellifer, b) Synodontis schoutedeni.
Achsen wie in Abb. 1.
Der Panzerwels Corydoras paleatus erzeugte Stridulationslaute auf folgende
Art und Weise: wurde der Hinterkörper beidseitig gefaßt, warf der Fisch seinen
Vorderkörper schnell hin und her. Dabei waren Laute hörbar, deren Sonagramme
Stridulationslauten ähneln (Abb.9). Wurde der Fisch am Rumpf und an einer Brust-
flosse festgehalten, blieb die andere Pectoralis arretiert. Im Gegensatz zur stridu-
lierenden Peckoltia zeigten die Loricariidae Ancistrus cf. dolichopterus, Panaque
nigrolineatus, Parancistrus aurantiacus und Pseudacanthicus spinosus aus derselben
Unterfamilie Ancistrinae zu keiner Tageszeit Stridulation. Alle Individuen dieser
178 ANDREAS HEYD & WOLFGANG PFEIFFER
Mi (1 , , sn e Ms fe tue
+ Aad igs È = 4 In
3 i Hal 88, Brit DIR br O A ri
ee eee
b è € i
1° | ih hs di L Fi; i |
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100 ms
Lu
| BA Bi
BR
| 4 | oti
Rie? 1 Duo
ABB. 5
Ausschnitt je eines Sonagramms von: a) Agamyxis flavopictus, b) Dysichthys coracoideus.
._ Achsen wie in Abb. I.
Species, ausgenommen Peckoltia, verhielten sich beim Herausnehmen aus dem
Wasser ähnlich. Sie spreizten und arretierten beide Brustflossen, richteten ihre
Rückentlosse auf und führten Schwanzschläge aus. Im Gegensatz zu Pseudacanthicus
klappten Ancistrus und Parancistrus zusätzlich ihre mit Hakenstacheln besetzten
Interopercula nach außen. Panaque besitzt keine solchen Interopercula. Anders als die
LAUTERZEUGUNG DER WELSE (SILUROIDEI) 179
ms
200
600
20 14
300 31
100 774010 19 32 2 44
ABB. 6
Dauer der Stridulationslaute (a) und der Pausen zwischen den Lauten (b) von 33 Individuen aus
11 Species. Dargestellt sind Mittelwert, Standardabweichung, Minimum und Maximum sowie
Anzahl der Laute (Ziffer). Abszisse: Species (Abkürzungen siehe Tab. 2; Pg = Pterygoplichthys
gibbiceps = Synonym von Glyptoperichthys gibbiceps = Gg). Ordinate: Dauer in ms. Beachte
die stärkere Dehnung der Ordinate in (a) gegenüber (b).
180 ANDREAS HEYD & WOLFGANG PFEIFFER
200
150
100
600
=
200 Za
BEER
As Bm Du Gg Pp Af Dc Mg Mv So Ss
ABB. 7
Box-whisker-plots der Lautdauer (a) und der Pausendauer (b) der 11 Species. Abszisse: Species
in Reihenfolge nach der Stridulationsweise: As-Pp Abduktion; Af-Ss Ab- und Adduktion. Ab-
kürzungen siehe Tab. 2. Ordinate: Dauer in ms. Box: Median, 25. und 75. Perzentil. Whisker:
10. und 90. Perzentil. Punkte: 5. und 95. Perzentil.
beiden Bagridae der Gattung Mystus stridulierten vier wiederholt gepriifte Exemplare
von Hemibagrus nemurus nicht, zeigten jedoch ein anderes bemerkenswertes Ver-
halten. Sie erzeugten mehrmals hintereinander durch Auspressen von Luft aus ihren
Kiemenspalten Quietschlaute von 26 - 56 ms Dauer, mit einer Hauptfrequenz von 1,5
kHz (Abb. 10). Bei Pimelodus pictus waren wenige Stridulationslaute gleichzeitig mit
Trommelmuskellauten zu hören.
LAUTERZEUGUNG DER WELSE (SILUROIDEI) 181
EE
As Bm Du Gg Pp Af Dc Mg Mv So Ss
ABB. 8
Verhaltnis von Lautdauer zu Pausendauer fiir jede Species, dargestellt als box-whisker-plots.
Erläuterung siehe Abb. 8.
TROMMELMECHANISMEN:
Pimelodus pictus: alle vier Individuen erzeugten Laute, die an das Summen
einer Fliege oder Hummel erinnern. Ein Exemplar stridulierte kurz außerhalb des
Wassers, wobei gleichzeitig das Summen ertönte. Auch ein im Becherglas stridu-
lierendes Exemplar summte. Die Summtöne waren etwa | sec lang und wurden mehr-
mals hintereinander erzeugt.
Arius seemani: beim Transport dieser Welse waren ebenso wie im Aquarium
Laute vernehmbar, die sich wie „Popps“ anhörten und entfernt an das Gackern von
Hühnern erinnerten. Auf Klopfen an das Aquarium stellten die Fische diese
Geräusche für einige Sekunden ein. Manche Individuen ließen diese Laute auch
außerhalb des Wassers hören, während sie gleichzeitig stridulierten. Diese Laute
waren leiser als das durch Stridulieren erzeugte Knarren.
ANDREAS HEYD & WOLFGANG PFEIFFER
182
[UT
RR iii
nee B
mL
BEA an fr
ARBRE tt 18:
100 ms
ABB. 9
on Corydoras paleatus (2 Exemplare). Achsen wie in Abb. 1.
le
Ausschnitte aus Sonaerammen v
LAUTERZEUGUNG DER WELSE (SILUROIDEI) 183
Li!
1
Fin
ye
Li
| |
À
{
hi
1
# E
=
= a
=. =
= at ae È
FA “= wu
a &
100 ms
ABB. 10
Zwei “Quietschlaute” eines Exemplars von Hemibagrus nemurus. a: Sonagramm. b: Frequenz-
analyse des Sonagraphen; vertikal Frequenz, horizontal Amplitude.
Versuch A: Die 4 Fische der ersten Gruppe begannen gleich nach dem
Einsetzen in das Versuchsaquarium Trommellaute zu erzeugen, die in | m Entfernung
vom Aquarium zu hören waren. Tagsüber lagen diese Welse entweder eng bei-
sammen ruhig am Boden oder ruhten mit dem Kopf nach oben fast senkrecht an einer
Aquarienwand. Bei Fütterung schwammen sie umher, danach ruhten sie wieder
beieinander. Auch die zweite Gruppe gab sofort nach dem Einsetzen ins Versuchs-
aquarium Trommellaute von sich. Sie verhielt sich insgesamt ruhiger als die erste
184 ANDREAS HEYD & WOLFGANG PFEIFFER
ABB. 11
24-Stunden-Lautaktivität der beiden Gruppen (Gr. 1, Gr. 2) von Arius seemani. Dargestellt sind
Medianwerte über 3 Tage. Abszisse: Uhrzeit. Ordinate: Anzahl der Laute.
Gruppe und erzeugte bei Fütterung und beim Hantieren am Aquarium weniger Laute.
Das Verhalten am Tag ähnelte dem der ersten Gruppe. Beide Gruppen zeigten in ihrer
Lautaktivität einen ausgeprägten Tag-Nacht-Rhythmus (Abb. 11). Die Lautproduktion
war am Tag deutlich höher als nachts und etwa | Stunde nach Einsetzen der
Lichtperiode am höchsten. Die Lautaktivität nahm dann ab, um gegen 12 Uhr ein
erneutes, aber kleineres Maximum zu erreichen. Am Nachmittag war ein weiteres,
schrittweises Abklingen der Lautanzahl zu verzeichnen. Zwischen 19 und 7 Uhr
(Dunkelphase) erzeugten die Welse nur wenige Laute. Bereits zwei Stunden vor der
Lichtperiode (7 Uhr) intensivierten beide Gruppen ihre Lautaktivität. Die Lautserien
waren am Morgen gehäuft, am Mittag vereinzelt und wurden oft nur von einem ein-
zigen Fisch erzeugt. Die nachts registrierten Laute waren dagegen meist Einzel-
geräusche, nur ausnahmsweise Serien. Bei Gruppe 1 war die Gesamtzahl der Laute in
der Hellphase um etwa 50 %, nachts um etwa 23 % höher als bei Gruppe 2.
Versuch B: Alle Laute des isolierten Exemplars zeigten gleiche Frequenz-
spektren. Die Hauptfrequenzen lagen unter 500 Hz. Die maximale Intensität war bei
140 - 200 Hz, hohe Intensitäten sind bei 250 - 400 Hz zu erkennen (Abb. 12). Die
mittlere Lautdauer betrug 209,5 + 17,4 ms, die Intervalldauer 0,3 - 6 sec.
LAUTERZEUGUNG DER WELSE (SILUROIDEI) 185
0 200 1000 ms
ABB. 12
Farbspektrogramm der Trommellaute von Arius seemani. Abszisse: Zeitachse in ms. Ordinate:
Frequenz in kHz. Farbskala rechts: Intensitätsverteilung des Schalldruckpegels in dB.
Struktur der Trommelmuskelapparate (Schwimmblasenmechanismen):
Arius seemani besitzt im Bereich der vorderen Schwimmblasenkammer, ein
laterales Fenster in der Muskulatur (“lateral cutaneous area”). Die Versteifung des
Schädel-Hals-Bereiches wird durch den mit dem Basioccipitale fest verwachsenen
Wirbelkomplex verstärkt. Eine knöcherne Lamelle des Epioticums ist mit dem
Transversalfortsatz des 4. Wirbels fest verbunden. Der Wirbelkomplex wird von den
Wirbeln 2 bis 7 gebildet. Die Wirbel 4 bis 7 formen mit ihren Transversalfortsätzen
eine der Schwimmblase aufliegende Platte. Während die Transversalfortsätze 6 und 7
nur starres Bindegewebe miteinander verbindet, sind die Fortsätze 4 und 5
miteinander verwachsen. Der anteriore Teil des Transversalfortsatzes des 4. Wirbels
ist zu einem langen, ventrad gebogenen Ast ausgezogen. Tavolga (1962) hat ihn zu
Ehren seines Entdeckers Johannes Müller (1842) ,,Miiller’ sche Ramus“ genannt
(Abb. 13). Dieser lanzenförmige Ramus liegt der Schwimmblase craniad auf und ist
mit ihr über Bindegewebe verwachsen, das ihn bis zum Ansatz des Trommelmuskels
überzieht. Dieser indirekte Trommelmechanismus wurde von Müller (1842) als
„Springfederapparat“ (,,elastic spring apparatus“) bezeichnet. Er wird vom Trans-
versalfortsatz des 4. Wirbels, dem Müller’schen Ramus und dem Trommelmuskel
(Protractor) gebildet. Der Protractor hat seinen Ursprung am Posttemporale, dem
Epioticum und dessen knöcherner Lamelle. Die Ansatzstellen des Muskels liegen am
Vorderrand des Transversalfortsatzes des 4. Wirbels bis zu dessen Verbindung mit
186 ANDREAS HEYD & WOLFGANG PFEIFFER
ABB. 13
Springfederapparat von Arius seemani (215 mm) der rechten Körperseite von lateral. P,
Protractor mit blutrotem anterioren Anteil. El, knöcherne Lamelle des Epioticums. TP,,
Transversalfortsatz des 4. Wirbels. MR, Müller’sche Ramus. 8:1.
der Epioticum-Lamelle. Ein weiterer Teil des Muskels zieht zum proximalen
Müller’schen Ramus. Der vordere Bereich des fächerförmigen Trommelmuskels
erscheint am frischen Präparat stärker blutrot als der zum Transversalfortsatz des 4.
Wirbels ziehende Ast. Beide Muskelbereiche lassen sich kaum voneinander trennen,
da sie gemeinsam den Protractor bilden (Abb. 13). Die herzförmige Schwimmblase
eines 260 mm großen Arius ıst 33 mm lang und erstreckt sich bis zum 8. Wirbel,
eingerahmt vom Müller’schen Ramus, dem Wirbelkomplex und dessen Transver-
salfortsätzen. Sie wird durch ein unvollständiges Diaphragma in eine anteriore und
eine posteriore Kammer geteilt. Letztere wird durch zwei Transversalsepten weiter
unterteilt. Die Ansatzstellen der Septen sind an der dorsalen und ventralen Schwimm-
blasenwand verbreitert (Abb. 14). Der kleine Ductus pneumaticus zieht vom caudalen
Ende der vorderen Schwimmblasenkammer zum Vorderdarm.
Pimelodus pictus: Die vordere Schwimmblasenkammer grenzt teilweise an die
Haut. Die Wirbelkörper 2 bis 5 sind miteinander verwachsen. Die Transver-
salfortsätze der Wirbel 4 und 5 bilden zusammen eine starre, der Schwimmblase auf-
liegende, leicht geschwungene Platte. Der Transversalfortsatz des 4. Wirbels teilt sich
distal in einen anterioren und einen posterioren Ast, an dessen wulstartig verstärktem
cranialem Rand der Trommelmuskel entspringt. Dieser Fortsatz ist gelenkig mit dem
LAUTERZEUGUNG DER WELSE (SILUROIDEI) 187
ABB. 14
Schwimmblase von Arius seemani (260 mm, männlich), linke Körperseite, lateral geöffnet. a)
anteriore Kammer, durch das Diaphragma (D) von der posterioren Kammer teilweise getrennt.
8:1. b) posteriore Kammer. Deutlich sichtbar die Transversalsepten (TS) und das Diaphragma
(D), das sich wie die TS and der dorsalen und ventralen Schwimmblasenwand verbreitert. 8:1.
188 ANDREAS HEYD & WOLFGANG PFEIFFER
ABB. 15
Schwimmblasen von ventral. a) Pimelodus pictus (105 mm). Der blutrote Trommelmuskel
(TM) überdeckt mehr als die Hälfte der Schwimmblase (SB). 8:1. b) Pimelodella gracilis
(123 mm). Schwimmblase frei, kein Trommelmuskel. 8:1.
LAUTERZEUGUNG DER WELSE (SILUROIDEI) 189
ABB. 16. Pimelodus pictus (Kopf nach links gelegen) a) Schwimmblase eines 105 mm langen
Exemplars von links lateral geöffnet. Die anteriore großlumige Kammer ist von der posterioren
Kammer durch ein Diaphragma (D) teilweise getrennt. Der hintere Schwimmblasenteil ist von
Transversalsepten (TS) weiter unterteilt. 12:1. b) Schwimmblase eines 120 mm langen for-
malin-fixierten Exemplars, von dorsal geöffnet. Deutlich sind die zur dorsalen Schwimm-
blasenwand ziehenden Septen, die sich an ihren Ansatzstellen verbreitern. 8:1.
190 ANDREAS HEYD & WOLFGANG PFEIFFER
Posttemporale verbunden. Der paarige, durch große Blutgefäße versorgte und daher
intensiv rote Trommelmuskel (Protractor) hat seinen Ursprung am Transversalfortsatz
des 4. Wirbels und reicht über die Hälfte der ventralen Schwimmblasenwand. Er
bedeckt die vordere Schwimmblasenkammer mit Ausnahme der “lateral cutaneous
areas” und setzt am cranialen Ende der hinteren Kammer an. An seiner Ansatzstelle
läuft der Trommelmuskel flach aus. Seine Ansatzareale sind S-förmig gekrümmt. Die
herzförmige Schwimmblase erstreckt sich vom 2. bis zum 6. Wirbel und ist bei einem
105 mm großen Exemplar 15 mm lang (Abb. 15). Sie wird durch ein unvollständiges
Diaphragma in eine anteriore und eine posteriore Kammer unterteilt. Weil das
Diaphragma von lateral-dorsal gelegenen Öffnungen durchbrochen wird, sind die
beiden Kammern miteinander verbunden. Die posteriore Kammer wird durch Trans-
versalsepten nur unvollständig weiter unterteilt. Die Septenbildung beginnt an der
Ansatzstelle des Trommelmuskels und reicht bis zum caudalen Schwimmblasenende.
Die Septen ziehen zur dorsalen und ventralen Schwimmblasenwand (Abb. 16). Der
Ductus pneumaticus entspringt in der Mitte des caudalen Endes der vorderen
Schwimmblasenkammer.
Pimelodella gracilis besitzt ebenso wie Pimelodus pictus und Arius seemani
in der lateralen Muskulatur beidseitig je ein ein Fenster, so daß ein Teil der vorderen
Schwimmblasenkammer, nur von Haut überdeckt, in engem Kontakt mit dem
Außenmedium Wasser steht. Die Wirbel 3 bis 5 sind miteinander verwachsen. Die
Transversalfortsätze der Wirbel 4 und 5 bilden zusammen eine der Schwimmblase
aufliegende Platte. Der anteriore Teil des Transversalfortsatzes des 4. Wirbels formt
an seinem cranialen Ende einen Wulst, ähnlich wie bei Pimelodus. Ein Trommel-
muskelapparat wurde bei Pimelodella jedoch nicht gefunden. Die herzförmige
Schwimmblase erstreckt sich vom 2. bis zum 6. Wirbel und ist bei einem 123 mm
großen Exemplar 8,5 mm lang. Sie wird von einem unvollständigen Diaphragma in
eine anteriore und eine posteriore Kammer unterteilt. Das Diaphragma besitzt rechts
und links je eine Öffnung, durch welche die beiden Kammern kommunizieren (Abb.
17). Die posteriore Kammer ist mindestens doppelt so lang wie die anteriore und wird
durch ein vertikales Längsseptum vollständig zweigeteilt. Weitere Transversalsepten
wurden, im Gegensatz zu Pimelodus, nicht gefunden. Der Ductus pneumaticus
entspringt an der Schnittstelle des Längsseptums mit dem Diaphragma.
LAUTERZEUGUNG UND SCHRECKREAKTION
In Tabelle 3 sind diejenigen Welsfamilien aufgelistet, die auf den Besitz von
Schreckstoffzellen und Schreckreaktion bzw. ihre Fähigkeit zur Lauterzeugung unter-
sucht worden sind. Ferner zeigt diese Übersicht das Vorkommen sowie die Anzahl
von Gattungen und Species nach Nelson (1994) und Teugels (1996). Der Reihenfolge
der Familien liegt ein Cladogramm von de Pinna (1998) zugrunde, wobei die
„niedrigen“ Taxa zuerst genannt sind. Während 20 der 23 dargestellten Taxa Schreck-
stoffzellen besitzen, ist die Schreckreaktion nur bei 5 Welsfamilien durch Video-
aufnahmen dokumentiert und bei 4 weiteren beobachtet worden. Stridulationsmecha-
nismen treten bei 14, Trommelmechanismen bei 8 Familien auf. Beide Lauter-
zeugungsmechanismen gemeinsam kommen bei 7 der „höheren“ Taxa vor.
LAUTERZEUGUNG DER WELSE (SILUROIDEI) 19]
ABB. 17. Pimelodella gracilis (Kopf nach rechts gelegen). a) Schwimmblase eines 123 mm
langen Exemplars, rechte Körperseite lateral geöffnet. Vordere und hintere Kammer sind durch
ein Diaphragma (D) teilweise getrennt. Die hintere Kammer wird von longitudinalem Septum
(LS) vollständig in zwei Hälften unterteilt. 12:1. b) Schwimmblase eines 100 mm Exemplars,
ventral geöffnet. Die seitlichen Öffnungen im Diaphragma sind im Vergleich zu Pimelodus
pictus großlumiger, weitere transversale Septen fehlen. 8:1.
192 ANDREAS HEYD & WOLFGANG PFEIFFER
TABELLE 3
Vorkommen, Anzahl der Gattungen (G.) und Species (Spec.) nach Nelson (1994) und Teugels
(1996) sowie das Auftreten von Schreckstoffzellen (SZ), Schreckreaktion (SR), Stridulation
(ST) und Trommelmechanismen (TM) bei verschiedenen Familien und Unterfamilien der
Welse. Reihenfolge der Familien nach einem Cladogramm in de Pinna 1998, p. 289;
Verwandtschaftsgruppen mit @ markiert. * : Unterfamilie der Pimelodidae = Rhamdiinae in
Teugels 1996. **: Familie bei de Pinna 1998 nicht aufgelistet.
Abkürzungen: Ab = Abduktion; Ad = Adduktion; AF = Afrika; AS = Asien; D = Rückenflosse;
E = Europa; MA = Mittelamerika; MAR = marin; NA = Nordamerika; PA = Pazifik; SA =
Südamerika; SW = Süßwasser; TS = tropisch-subtropisch; V = Videoaufnahmen von SR vor-
handen; + = vorhanden; (+) = SR gesehen; + ? = vorhanden, Mechanismus nicht beschrieben; ?
= fraglich; -- = nicht vorhanden.
G./ Spec.
G./Spec. SZ SR ST ™
Familie Vor-
kommen Nelson 1994 Teugels 1996
@ Cetopsidae SW/SA 4/12 4/14 +
@ Loricariidae SW /SA, MA 80 / > 550 88/651 -- -- Ab
Callichthyidae SW /SA, MA 71130 7/144 + V Ab
Trichomycteridae SW / MA, SA 36 / 155 40/154 +
Amphiliidae SW / AF 7/47 9/60 +
Sisoridae SW/AS 20/85 20/98 + D
@ Aspredinidae SW/SA 10 / 32 10 / 32 -- -- Ab+Ad
© Heptapterinae* SW/SA : 18 / 220 Ab
@ Ictaluridae SW/NA 7145 7145 + (+) Ab/Ab+Ad
@ Doradidae SW/SA 35 / 90 37 / 94 + Ab+Ad +
Auchenipteridae SW / SA 21/60 20/61 + + +
Mochokidae SW / AF 10 / 167 11/177 + (+) Ab+Ad +
Ariidae MAR (SW) / TS 14 / 120 14/148 + Ab +
@ Siluridae SW/E, AS 12/100 9/60 + V
Malapteruridae SW / AF 12 10/73 + +
Chacidae SW / AS 193 13 +
Plotosidae MAR, SW / AS, PA 9/32 932 + V ?
Clariidae SW / AF, AS 13 / 100 15 / 92 Et (Gr) ?
@ Pangasiidae SW / AS 2/21 2/2] + V + +
Schilbeidae SW / AF, AS 18/45 13751 + V
@ Bagridae SW / AF, AS 30/210 [Sys SEE Ab+Ad +
@ Pimelodinae SW / MA, SA 56 / 300 931/323 ab (©) Ab +
@ Heteropneustidae** SW / AS 17/22 YE + apa
DISKUSSION
LAUTERZEUGUNG:
Ariidae: Die Unterwasserlaute von Arius felis (Synonym Galeichthys felis) stu-
dierte Dobrin (1947). Diese Species erzeugt sowohl Laute mit dem Springfeder-
apparat (Burkenroad, 1931) als auch Stridulationsgeräusche. Frisch gefangene A. felis
brachten bei Bewegung der Brustflossen 30 - 50 ms lange Laute im Frequenzbereich
2 - 4 kHz hervor (Tavolga, 1960). Arius seemani produziert bei Abduktion der
Brustflossen Stridulationslaute, deren Dauer und Frequenzanteile ebenfalls in diesem
Bereich liegen. Die Lauterzeugung mittels Springfederapparat wurde von Tavolga
LAUTERZEUGUNG DER WELSE (SILUROIDEI) 193
(1971b, 1976, 1977) eingehend untersucht. Arius felis erzeugt fast dauernd Trom-
mellaute der Frequenz 100 Hz und benützt sie als primitives Echolot bis auf 10 cm
Entfernung. Bemerkenswert ist die zu dieser tiefen Frequenz passende Hörschärfe.
Arius felis hört im Bereich 50 - 1000 Hz, mit bester Hörschärfe bei 100 - 200 Hz
(Popper & Tavolga, 1981). Diese Werte liegen weit unter denen von Ameiurus
nebulosus (Ictaluridae) mit 100 - 4000 Hz bzw. 600 - 800 Hz (Poggendorf, 1952,
Weiss et al., 1969). In Feldstudien wurde gezeigt, daß A. felis besonders nachts Laute
ausstößt, die dem Zusammenhalt der Gruppe dienen sollen. In Laborversuchen waren
die Laute in Gesellschaft kürzer als unter Isolation. Ungestört gaben die Welse leise,
20 - 40 ms kurze Grunzer von sich; bei Störung dagegen wurden laute, 100 - 150 ms
lange Geräusche mit einer Grundfrequenz von 150 Hz registriert (Tavolga, 1960,
1976). Bei Langzeituntersuchungen an A. felis fand Breder (1968), daß die Welse von
April bis Oktober mit einer Sommerpause im Juli/August Laute erzeugten, haupt-
sächlich von 17 - 23 Uhr bei einer Wassertemperatur von 23 - 31°C. Die Trommel-
aktivität von A. seemani zeigte im Labor einen anderen Tag-Nacht-Rhythmus. Sie
begann vor der Lichtperiode und war in den Morgenstunden am höchsten; nachts
wurden kaum Laute registriert. Eine Funktion der Laute für den Zusammenhalt der
Gruppe oder für die Orientierung im Dunkeln ist daher unwahrscheinlich. Bei
Fischarten außerhalb der Welse wurden ebenfalls Dämmerungslaute (morgens und
abends) beschrieben, deren biologische Bedeutung jedoch unbekannt ist (Winn, 1964;
Mehickvet al 11999).
Im Gegensatz zu den Untersuchungen an A. felis (Tavolga 1960) unterbrach A.
seemani seine Trommelmuskelaktivität bei Störungen kurz, und die Lautdauer (ca.
200 ms) blieb vor und nach Störungen nahezu unverändert. Die Grundfrequenz der
Laute lag mit 140 - 200 Hz ım Bereich der von A. felis bekannten Kontraktions-
frequenz des Trommelmuskels (Fusionsgrenze < 400 Hz). Die anatomischen Über-
einstimmungen zwischen beiden Species deuten darauf hin, daß auch bei A. seemani
die Grundfrequenz der Laute der Kontraktionsfrequenz des Protractors entspricht. Die
Pausen zwischen den Lauten eines isolierten A. seemani schwankten in Ruhe und bei
Störung. Arius seemani besitzt einen Springfederapparat wie A. felis (Tavolga 1962).
Einen solchen Apparat führen auch die Auchenipteridae, Bagridae, Doradidae,
Malapteruridae, Mochokidae und Pangasiidae (Müller, 1842; Bridge & Haddon,
1893; Chranilov, 1929; Tavolga, 1962). Sörensen (1884) sowie Bridge & Haddon
(1894) wiesen bereits darauf hin, daß der Springfederapparat der Doradidae der Laut-
erzeugung dient, wie von Kastberger (1977, 1978) bestätigt. Entsprechendes gilt für
die Mochokidae (Abu-Gideiri & Nasr, 1973). Der Springfederapparat und der direkte
Trommelmechanismus (bei Pimelodidae) haben sich möglicherweise bei verschie-
denen Welsfamilien unabhängig voneinander entwickelt (Alexander, 1965). Der
Funktionsmechanismus des Springfederapparats läßt sich folgendermaßen beschrei-
ben: Durch Kontraktion des Trommelmuskels (Protractors) werden der „Müller’sche
Ramus“ (Tavolga, 1962) und der Transversalfortsatz des 4. Wirbels (Knochenplatte)
craniad bzw. dorsad bewegt. Weil der Müller’sche Ramus über Bindegewebe mit der
Tunica externa der vorderen Schwimmblasenkammer verwachsen ist, wird die
Schwimmblase bei seiner Bewegung mitgezogen. Entspannt sich der Protractor,
194 ANDREAS HEYD & WOLFGANG PFEIFFER
schnellen Knochenplatte und Müller’scher Ramus wieder zurück. Die vordere
Schwimmblasenkammer wird komprimiert und ihr Gasinhalt in die hintere Kammer
gepreBt. Die Schwimmblase ist durch Transversalsepten unterteilt. Das an den
Septenrändern und dem Diaphragma vorbeiströmende Gas versetzt diese in Schwin-
gung. Die Schwimmblasenwand wird zum Vibrieren angeregt und ihre Schwingung
durch die trommelnden Knochenelemente verstärkt (vergleichbar einem Schlagzeug).
Die Schwimmblase dient als Resonanzkörper (Trommel). Schwingungsgeneratoren
sind der Trommelmuskel und die knöchernen Elemente. Die Synchronisation der
beiden Trommelmuskeln, rechts und links am jeweiligen Müller’schen Ramus, wird
zurückgeführt auf die polyaxonale und multiple Versorgung ihrer Fasern durch den
zweiten Spinalnerven (Schneider, 1967).
Aspredinidae: Die Stridulationslaute von Bunocephalus spec. wurden von
Winn (1964) analysiert und von Pfeiffer & Eisenberg (1965) an einigen Species
gehört. Die Kontraktionsdauer der Muskelpotentiale von Bunocephalus spec. ent-
spricht mit 100 ms der Lautlänge (Gainer, 1967). Die Lautdauer von Dysichthys
coracoideus ist mit 80 ms nur wenig kürzer. Wie Bunocephalus spec. striduliert D.
coracoideus bei Abduktion und Adduktion.
Auchenipteridae: Sie besitzen neben der Brustflossenstridulation auch einen
Springfederapparat (Müller 1842). Die Trommellaute von Trachycorystes spec. haben
eine Grundfrequenz von 120 Hz und können einige Sekunden andauern (Kastberger,
1978). Weitere Daten zu Schwimmblasen- und Stridulationslauten finden sich bei
Kaatz (1999).
Bagridae: Leiocassis micropogon erzeugt „knurrende“, L. siamensis und L.
poecilopterus produzieren „krächzende“ Laute. L. brashnikowi gibt „einzelne Töne“
von sich, und beim Paarungsspiel von Mystus vittatus entstehen „zwitschernde“ Laute
(Riehl & Baensch, 1983; Franke, 1985; Sterba, 1990), wobei der Mechanismus der
Lauterzeugung nicht genannt wird. Mystus vittatus und M. gulio stridulieren bei
Abduktion und Adduktion der Brustflossen. Mit den „Quietschlauten“ von Hemi-
bagrus nemurus gibt es eine weitere, nämlich pneumatische Weise der Lautent-
stehung. Agassız (1852) hörte bei einem Wels Geräusche, die anscheinend durch
Luftausstossen aus der Schwimmblase hervorgerufen wurden. Bei Synodontis schall
ist ein „Atemgeräusch“ vernehmbar, das vermutlich mit der Schwimmblase zusam-
menhänst (Dufossé 1874). Sörensen (1895) beschrieb ein katzenähnliches Fauchen
(„not unlike the hissing of a cat”) beim Zitterwels Malapterurus und führte dies auf
die zwischen hinterer und vorderer Schwimmblasenkammer durch einen dünnen
Gang streichende Luft zurück. Kaatz (1999) beobachtete Lauterzeugung durch
Luftausstoßen bei 8 Welsfamilien.
Callichthyidae: Callichthys callichthys läßt bei der Brutpflege „grunzende“
Laute hören (Riehl & Baensch, 1983). Hoplosternum thoracatum erzeugt bei
Geschlechtsreife im Wasser Stridulationslaute (Mayr 1987). Für Dianema urostriata
wurden Stridulationslaute nur bei Abduktion nachgewiesen, wie von H. thoracatum
bekannt. Bei Corydoras paleatus stridulieren beide Geschlechter beim Fang, die
Männchen auch bei Balz und Paarung (Pruzsinsky & Ladich, 1998).
LAUTERZEUGUNG DER WELSE (SILUROIDEI) 195
Clariidae / Heteropneustidae: Stridulationsgeräusche wurden für Hetero-
pneustes fossilis beschrieben. Die Laute entstehen zum einen durch Bewegung der
Brustflossen, zum anderen durch Reiben der Pharyngealzähne am Mundboden. Es
wird vermutet, daß die Laute der Warnung und Abwehr von Angreifern sowie
während der Fortpflanzungsperiode der Schwarmbildung dienen (Agrawal & Sharma
1965). Diese Species, früher zu den Clariidae gezählt, wird heute den Hete-
ropneustidae zugeordnet (Burgess, 1989; Teugels, 1996). Über die Lauterzeugung der
Clariidae gibt es kaum Zeugnisse: Tennent (1859; zitiert in Sörensen, 1895, p. 115f.)
berichtet über Aussagen von Fischern, nach denen ein „Magoora“ genannter Fisch in
einem See nahe Colombo bei Störung im Wasser „Grunzlaute‘“ hervorbringt. Nach
Day (1881) handelt es sich dabei um einen Wels der Gattung Clarias, wahrscheinlich
um Clarias magur (,,Mah-gur* oder „Magurah“, Day, 1958), einem Synonym von
Clarias batrachus (Eschmeyer, 1998). Der Laut soll durch Schwimmblasenluft ent-
stehen, die über den Ductus pneumaticus entweicht (Sörensen, 1895). Es könnte sich
aber auch um Stridulationslaute handeln, da viele Clariidae kräftige Brustflossen-
stacheln besitzen, deren Struktur derjenigen stridulierender Species ähnelt (Nawar
1954).
Doradidae: Acanthodoras spinosissimus „grunzt“ (Innes 1956), „Knurrt‘ (Ster-
ba 1956), „krächzt‘ und gibt einen lang anhaltenden Quarrlaut von sich, der aus einer
Folge von Einzellauten besteht (Villwock, 1960). Amblydoras hancocki erzeugt
knurrende Töne, und Platydoras costatus stößt beim Fangen „quakende“ Laute aus
(Sterba 1990). Die Lauterzeugung in Abhängigkeit von der Bewegung der Brust-
flossen wurde erstmals von Pfeiffer & Eisenberg (1965) experimentell untersucht,
durch gleichzeitige Film- und Tonbandaufnahmen von Agamyxis albomaculatus,
Amblydoras hancocki und Platydoras costatus. Die Laute entstehen bei Abduktion
und Adduktion, wobei die Abduktionslaute länger sind als die Adduktionslaute. Die
an Agamyxis flavopictus erhobenen Befunde bezüglich Laut- und Pausenlängen
entsprechen den Ergebnissen von Pfeiffer & Eisenberg (1965). Dagegen fand Ladich
(1997), daß sich Ab- und Adduktionslaute bei Agamyxis pectinifrons bzw. Platydoras
costatus ın der Länge nicht unterscheiden. Die Dauer der Stridulationslaute liegt bei
den beiden Species A. flavopictus und A. pectinifrons mit ca. 100 ms im gleichen
Bereich. A. flavopictus wird 1m Handel oft als A. pectinifrons bezeichnet (Franke
1985). Hassar orestis kann ebenfalls stridulieren und ist durch seine abgespreizten
Flossenstacheln sogar vor Piranhas geschützt (Markl, 1968). Neben der Brustflossen-
stridulation besitzen die Doradidae einen indirekten Trommelmechanismus, den
“Springfederapparat” (Müller, 1842; Sörensen, 1884, 1895; Bridge & Haddon, 1894;
Dorn, 1976; Kastberger, 1977, 1978). Die biologische Bedeutung der durch die
Brustflossenstridulation erzeugten Laute sehen Pfeiffer & Eisenberg (1965) im
„Schutz oder der Verteidigung“, in der Warnung eines potentiellen Beutegreifers vor
einem nur schwer zu verschlingenden, gefährlichen Happen. Die Gefährlichkeit ist
gegeben durch die Sperrmechanismen der Stacheln von Brustflossen und Rücken-
flosse (Brousseau, 1976), in Verbindung mit ihrer großen mechanischen Festigkeit
(Schaefer, 1984) und möglichen Giftigkeit (Birkhead, 1972), die sogar für den
Menschen tödlich sein kann (Nelson, 1994).
196 ANDREAS HEYD & WOLFGANG PFEIFFER
Ictaluridae: Den Bau des Schultergürtels von /ctalurus nebulosus beschreibt
Brousseau (1976). I. nebulosus striduliert bei Abduktion und Adduktion, wodurch die
Anzahl der aggressiven Auseinandersetzungen zwischen dem Revierinhaber und
einem Eindringling herabgesetzt wird (Rigley & Muir, 1979). Dagegen striduliert
Ictalurus punctatus nur bei Abduktion, meistens mit der rechten Brustflosse (Fine et
al., 1996, 1997). Daß /ctalurus nebulosus nach Lundberg (1992) zur Gattung
Ameiurus gehört, könnte den Unterschied im Stridulationsmechanismus erklären.
Jedoch sind nach Eschmeyer (1998) beide Gattungsnamen für diese Species gültig.
Die Sperrmechanismen des Rückenflossenstachels und der Brustflossenstacheln
schützen /. melas weitgehend davor, von Reihern und Rothalstauchern gefressen zu
werden, die ihre Beute nicht zerteilen, nicht aber vor Seeadlern, die sie zerlegen. Die
beiden kleineren Vogelarten fressen überwiegend Barsche (Forbes, 1989).
Loricariidae: Plecostomus spec. striduliert und besitzt ein ähnliches Brust-
flossenskelett wie die Doradidae und Mochokidae (Pfeiffer & Eisenberg, 1965).
Peckoltia pulcher und Glyptoperichthys gibbiceps stridulieren nur bei Abduktion. Bei
Pterygoplichthys spec. beträgt die Lautdauer 250 ms, die Grundfrequenz 500 Hz, die
Pausendauer 400 - 600 ms. Große Exemplare zeigen eine kürzere Laut- und Pausen-
dauer und ein höher liegendes Frequenzspektrum als kleine (Schachner, 1977).
Flossenstacheln und Schultergürtel von Prerygoplichthys spec. sind von enormer
mechanischer Stabilität (Schaefer, 1984).
Mochokidae: Synodontis spec. ist bereits vor 170 Jahren als erster durch
Stridulation Laute erzeugender Wels in die Literatur eingegangen (Geoffroy Saint-
Hilaire, 1829). Aus Daget et al. (1986) geht hervor, daß es sich hierbei um Synodontis
schall (Bloch & Schneider, 1801) handelt. Diese Species striduliert bei Abduktion
und Adduktion der Brustflosse (Müller, 1857, Dufossé, 1874) wie S. nigrita und S.
nigriventris (Pfeiffer & Eisenberg, 1965), S. ocellifer und S. schoutedeni. Die Be-
funde an S. ocellifer und S. schoutedeni entsprechen in jeder Beziehung (Lautmecha-
nismus, Laut- und Pausendauer, Struktur der Sonagramme und Frequenzschwer-
punkte) den vor 35 Jahre vom Zweit-Autor erhobenen (Pfeiffer & Eisenberg, 1965).
Alle gepriiften Mochokidae stridulieren schneller als die Doradidae, weshalb die
Dauer ihrer Laute und Intervalle kürzer ist. Ladich (1997) konnte keinen signifikanten
Unterschied in den Lautlängen zwischen diesen beiden Familien finden, was für die
Abduktionslaute knapp zutrifft (p = 0.08). Die Adduktionslaute der Doradidae waren
länger als die der Mochokidae. Zur Klärung dieser Frage sollten weitere Species
beider Familien untersucht werden. Die Mochokidae besitzen außer der Brustflossen-
Stridulation auch einen Springfederapparat (Sörensen, 1884, 1895; Bridge & Haddon,
1893). S. schall striduliert und trommelt. Er soll Laute erzeugen bei Verfolgung,
„Schmerz“, aggressivem Verhalten, Laichverhalten und als Reaktion auf Berührungs-
reize (Abu-Gideiri & Nasr, 1973).
Pangasiidae: Pangasius sutchi erzeugt Stridulationsgeräusche (Kaatz, 1999).
Pimelodidae: Bau und Funktion des Stridulationsapparates und des Trommel-
mechanismus wurden von Schachner (1977) an Pimelodus spec. untersucht. Schach-
ner & Schaller (1982) zeigen, daß es sich dabei um Rhamdia sebae handelt und
beschreiben Defensivgeräusche (Stridulation), Drohlaute (Intervalle 60 - 70 ms) und
LAUTERZEUGUNG DER WELSE (SILUROIDEI) 197
„Störungslaute“ (Trommellaute wie die Drohlaute, jedoch mit kürzeren Intervallen
von nur 10 - 20 ms und höherer Frequenz). Die Stridulationsgeräusche dauern 40 - 85
ms, im Mittel 54 ms; ihr Energieschwerpunkt liegt bei 2 KHz (Schachner 1977). Die
Stridulationsgeräusche von Brachyrhamdia meesi entstehen nur bei Abduktion der
Brustflosse, wie diejenigen von R. sebae. Auch bei dem von Kratochvil er al. (1980)
und Kratochvil & Völlenkle (1981) untersuchten Pimelodus spec. dienen die
Stridulationslaute bei Rivalenkämpfen als Warnlaute, zur Revierverteidigung und bei
interspecifischen Attacken. Stoß und Biß werden mit Stridulationsgeräuschen beant-
wortet. Akustische Drohsignale können Fluchtreaktionen auslösen. Die Trommellaute
dienen der innerartlichen Verteidigung, die Stridulationslaute der Warnung. Den
Feinbau der Trommelmuskeln beschreibt Dorn (1976). Die für die Lauterzeugung
verantwortlichen Motoneuronen identifizierten Ladich & Fine (1994) an Pimelodus
blochi und P. pictus. Drei Äste des N. occipitalis und die beiden ersten Spinalnerven
versorgen das Trommelorgan, ein rostraler Ast des N. occipitalis und die beiden
ersten Spinalnerven die Stridulationsmuskeln (Abduktor und Adduktor). Pimelodus
pictus zeigte Drohverhalten gegenüber Artgenossen durch Spreizen seiner Flossen
und Kiemendeckel, gleichzeitig Trommellaute erzeugend, wie von Schachner &
Schaller (1982) fiir Rhamdia sebae geschildert. Auch die Strukturen des Trommel-
mechanismus von P. pictus entsprechen weitgehend der Beschreibung für R. sebae. In
beiden Fällen handelt es sich um einen direkten Trommelmuskelapparat. Der
anteriore Teil des transversalen Fortsatzes vom 4. Wirbel bildet einen Wulst, an dem
der Trommelmuskel ventral entspringt. Bei beiden Species setzt der Trommelmuskel
S-förmig an der ventralen Schwimmblasenwand an, wobei er bei P. pictus etwas
weiter tiber den caudalen Rand der vorderen Schwimmblasenkammer hinwegzieht als
bei R. sebae. Die Kammerung des posterioren Schwimmblasenteils ist verschieden:
bei R. sebae treten zahlreiche Transversalsepten auf, bei P. pictus nur drei. Laut-
bildung und Funktion der Transversalsepten einschließlich des Diaphragmas der
Schwimmblase wurden bereits von Sörensen (1884, 1895) beschrieben. Durch
Kontraktion des Trommelmuskels wird die vordere Schwimmblasenkammer kompri-
miert und ihr Gasinhalt in die hintere Kammer gepreßt. Beim Entspannen des
Muskels wird das Gas wieder zurückgesogen. An den Rändern der unvollständigen
Transversalsepten vorbeistreichend versetzt es diese in Vibration und regt die gesamte
Schwimmblase zum Schwingen an. Der kontraktierende Trommelmuskel erzeugt das
Geräusch, das durch die Schwingungen von Septen und Schwimmblase verstärkt
wird. Sörensen (1895) schnitt winzige Löcher in die Schwimmblasenwand, wodurch
der Laut an Intensität verlor. Doch war der Laut sogar nach Entfernen der Schwimm-
blase noch schwach zu hören. Dagegen war nach Durchtrennung des paarigen Trom-
melmuskels kein Geräusch mehr wahrzunehmen. Bei R. sebae sind im posterioren
Schwimmblasenabschnitt nur die Septen beweglich, die Schwimmblasenwand ist starr
(Schachner & Schaller, 1982). Außer den Septen wird vor allem der vordere
Schwimmblasenabschnitt zum Schwingen angeregt, so daß die Vibrationen beidseitig
über die lateralen Fenster in der Muskulatur direkt an das umgebende Wasser
abgegeben werden können. Bei der Präparation der Schwimmblase von P. pictus war
auffallend, daß ihr posteriorer Teil stabiler ist als ihr anteriorer. Wurde die
198 ANDREAS HEYD & WOLFGANG PFEIFFER
Schwimmblase lateral eröffnet, fiel die vordere Kammer zusammen, während die
hintere stabil blieb. Der posteriore Schwimmblasenteil wird durch die Transver-
salsepten und die festere Wand gestützt. Tavolga (1962) reizte Trommelmuskeln von
Bagre marinus elektrisch: die zur Dauerkontraktion notwendige Frequenz beträgt 500
Hz. Die Fasern der Trommelmuskeln von P. pictus gehören zu den schnell zuckenden
und sind dicht kapillarisiert. Ontogenetisch leiten sich die Trommelmuskeln von
Myotom-Muskulatur ab, die in Richtung Peritoneum gewandert ist und sich diesem
aufgelagert hat. Das Peritoneum ist an der Ansatzstelle des Trommelmuskels an der
Schwimmblase mit deren Tunica externa verwachsen (Alexander, 1965). Der direkte
Trommelmuskelapparat deutet darauf hin, daß P. pictus Trommellaute erzeugt.
Pimelodella gracilis besitzt wie Pimelodus pictus beidseits je ein Fenster in der
Muskulatur, Transversalfortsätze des 4. und 5. Wirbels und eine gut ausgebildete
Schwimmblase. Ein Trommelmuskel konnte bei P. gracilis nicht gefunden werden.
Da nur juvenile Weibchen untersucht wurden, ist nicht auszuschließen, daß der
Trommelmuskel bei diesen Fischen entweder noch nicht entwickelt war oder nur bei
Männchen existiert. Die folgenden Tatsachen sprechen gegen einen Trommelapparat
bei P. gracilis: ein derart mächtiger Trommelmuskel, wie ihn P. pictus und Arius
seemani haben, wird vermutlich bereits in einem frühen Entwicklungsstadium an-
gelegt; die untersuchten juvenilen P. pictus besaßen bereits einen Trommelmecha-
nismus. Ferner spricht die Form der Schwimmblase von P. gracilis gegen eine
Funktion als Lautorgan. Schließlich fehlen der hinteren Schwimmblasenkammer
Transversalsepten, die nach Sörensen (1895) für die Lauterzeugung notwendig sind.
Die posteriore Kammer ist durch ein Longitudinalseptum zweigeteilt. Das Gas in der
Schwimmblase hat nur die Möglichkeit, an den Diaphragma-Rändern entlang in eine
dieser beiden hinteren Kammern zu gelangen. Zudem ist die vordere Kammer von
P. gracilis die kleinere, im Gegensatz zu P. pictus. Es könnte also nur ein geringes
Gasvolumen bewegt werden, um das Diaphragma und die Schwimmblasenwände
zum Schwingen anzuregen. Die beiden lateralen Fenster in der Muskulatur und die
knöchernen Elemente des 4. und 5. Wirbels sind für das Hören wichtig (Alexander,
1965). Diese Fenster verringern den Widerstand der Körperwand, so daß die
Schwimmblase Schallschwingungen fast ungedämpft aufnehmen kann, um sie über
die ihr craniad-dorsad aufliegenden Weberschen Knöchelchen an das Innenohr
weiterzuleiten.
Plotosidae: Burgess (1989) zitiert Beschreibungen von Plotosus lineatus, in
denen diese Species auch ,,bumblebee catfish“ (Hummelwels) genannt wird. Dem-
nach soll ein Summen hörbar sein, sobald man diese Welse aus dem Wasser fischt.
Auch bei Störungen im Wasser wurden solche Geräusche vernommen. Da ein
Schwimmblasenmechanismus bei dieser Species nicht beschrieben wurde, könnte es
sich um Stridulationslaute handeln.
Siluridae: Stridulationsgeräusche wurden für diese Familie bisher nicht
beschrieben. Bei Hawkins (1986) findet sich aber folgende Bemerkung: „Some
catfish of the family Siluridae produce a squeak when the enlarged pectoral spines are
moved.“ Da eine Quellenangabe fehlt, handelt es sich möglicherweise um eine
Verwechslung mit anderen Welsfamilien.
LAUTERZEUGUNG DER WELSE (SILUROIDEI) 199
Sisoridae: Im Gegensatz zu allen anderen Welsfamilien stridulieren die beiden
bisher untersuchten Sisoridae nicht mit den Brustflossen, sondern mit der Rücken-
flosse, wie von Haddon (1881) an Gagata gagata entdeckt und von Mahajan (1963)
an Sisor rhabdophorus bestätigt. Die Lauterzeugung geschieht bei Abduktion und
Adduktion der Rückenflosse und dient vermutlich dazu, Beutegreifer vor der Gefähr-
lichkeit oder Ungenießbarkeit des Fisches wegen seiner kräftigen Brustflossen-
stacheln und seines Rückenflossenstachels zu warnen.
Trommelmechanismen sind nach der Stridulation die häufigste Form der
Lauterzeugung. Die Frage, ob die Schwimmblase für die Lauterzeugung notwendig
ist, wird unterschiedlich beantwortet, obwohl es auch lauterzeugende Fische ohne
Schwimmblase gibt. Während die Schwimmblase schon früh als hydrostatisches
Organ erkannt wurde (Boyle, 1670, 1675; Ray, 1675), ist ihre große Bedeutung für
Hörvermögen, Weberschen Apparat, Lauterzeugung und akustische Kommunikation
erst seit dem 19. und 20. Jahrhundert bekannt: (1) Hörvermögen (anatomisch):
Weber, 1820; Reissner, 1859; Bridge & Haddon, 1893; Chranilov, 1929 und Chardon,
1968; (physiologisch) von Frisch & Stetter, 1932; Poggendorf, 1952 und Weiss et a/.,
1969; (Reviews) Harden Jones & Marshall, 1953; Alexander, 1964, 1965, 1966;
Popper & Fay, 1973, 1993; Rogers et al., 1988 und Schellart & Wubbels, 1998; (2)
Lauterzeugung (anatomisch): Müller, 1857; Dufossé, 1874; Moreau, 1876; Sörensen,
1884, 1895 und Bridge & Haddon, 1889; (Reviews) Harden Jones & Marshall, 1953;
Schneider, 1961, 1967 und Demski, et al., 1973; (3) akustische Kommunikation
(Reviews): Tavolga, 1960, 1971a, 1977, Fine, et al., 1977; Myrberg, 1981; Hawkins
& Myrberg, 1983; Michelsen, 1983; Urick, 1983 und Tyack, 1998.
Die Schwimmblase ist bei der Lauterzeugung durch Trommelmechanismen
der Siluroidei beteiligt (Miiller, 1857; Tavolga, 1962; Ladich, 1997). Durch ihre
Gasfillung fungiert sie als Impedanzwandler zwischen den Trommelmuskeln und
dem Wasser und verstärkt so die erzeugten Schwingungen. Es wird diskutiert, ob
dabei diejenigen Frequenzen am besten verstärkt werden, die ihrer Resonanzfrequenz
entsprechen (Demski ef al., 1973). Kaatz (1995) fand bei mehreren Species der
Auchenipteridae und Doradidae Korrelationen zwischen der Schwimmblasengröße
und der Hauptfrequenz der erzeugten Laute. Bei Fischarten außerhalb der Siluroidei
wurde festgestellt, daß größere Exemplare tiefere Laute hervorbringen. Dies wird auf
veränderte Resonanzeigenschaften der Schwimmblase zurückgeführt (Demski et al.,
1973), da die Resonanzfrequenz mit zunehmendem Volumen der Schwimmblase
sinkt (Urick, 1983). Es muß dabei berücksichtigt werden, daß die Schwimmblase von
verschiedenen Geweben umgeben ist, die Schwingungsdämpfungen bewirken und
ihre Resonanzeigenschaften beeinflussen. Bei Stridulationsgeräuschen ist die Betei-
ligung der Schwimmblase umstritten: in der Lautstärke besteht außerhalb des Wassers
bei intakten Doradidae und Mochokidae einerseits und Exemplaren mit eröffneter
Schwimmblase andererseits kein Unterschied (Pfeiffer & Eisenberg, 1965). Zur
selben Ansicht kamen Fine er al. (1997) bei Untersuchungen an /ctalurus punctatus:
hier spielt die Schwimmblase bezüglich Frequenzspektrum und Amplitude der Stridu-
lationslaute zumindest keine Hauptrolle, während eher der Schultergürtel auf die
200 ANDREAS HEYD & WOLFGANG PFEIFFER
Frequenzverteilung Einfluß nimmt. Andererseits ergaben Untersuchungen an Cory-
doras paleatus, daß die Hauptfrequenz der Stridulationslaute mit zunehmender
Körpermasse abnimmt (Pruzsinsky & Ladich 1998). Es wäre möglich, daß hier
veränderte Resonanzeigenschaften der Schwimmblase eine Rolle spielen. Ob stridu-
lierende Fische die Resonanzfrequenz ihrer Schwimmblase modulieren, wurde bisher
nicht untersucht (Zelick er al., 1999).
LAUTERZEUGUNG UND PHYLOGENIE
Wichtige Fragen zu den Welsen (Berg, 1958; Nelson, 1976; Burgess, 1989)
sind ungelöst. Über die Phylogenie und das Natürliche System gibt es verschiedene,
zum Teil konträre Meinungen (Eigenmann & Eigenmann, 1890; Regan, 1911, Green-
wood et al., 1966; Rosen & Greenwood, 1970; Roberts, 1973; Lauder & Liem, 1983;
Fink & Fink, 1981, 1996; Schaefer, 1987). Die Klassifizierung in Familien sowie die
Anzahl der Gattungen und Species hat sich laufend verändert (Tab. 3; Mo, 1991;
Nelson, 1994; Teugels, 1996). Auch die Verbreitungsgeschichte der Welse ist unklar:
diskutiert werden primäre Zentren in Südamerika und Südostasien, ein sekundäres
Zentrum in Afrika (Gosline, 1975) oder Entstehung in Südamerika bzw. Südostasien
(Novacek & Marshall, 1976; Briggs, 1979). Die Fossilgeschichte ist ebenfalls
unbefriedigend; vermutlich war die interkontinentale Verbreitung schon vor dem
Eozän abgeschlossen (Gosline, 1975). Innerhalb der Siluroidei gibt es so viel Parallel-
Evolution, daß die phylogenetischen Zusammenhänge bei unserem gegenwärtigen
Wissensstand unklar bleiben (Alexander, 1965; Teugels, 1996; de Pinna, 1998).
In ihrer Kommunikation zeigen die Welse ein vielfältiges Bild. Sie verfügen
über mindestens 6 verschiedene Möglichkeiten der Lauterzeugung (Tab. 3): (1)
Stridulation durch Abduktion der Brustflossen (Ariidae, Callichthyidae, Ictaluridae,
Loricariidae, Pimelodidae), (2) Stridulation durch Abduktion und Adduktion der
Brustflossen (Aspredinidae, Bagridae, Doradidae, Ictaluridae, Mochokidae), (3) Stri-
dulation mit der Rückenflosse (Sisoridae), (4) pneumatische Mechanismen durch
Luftausstoßen über die Kiemenspalten (Hemibagrus nemurus), (5) direkter Trommel-
mechanismus (Pimelodus pictus) und (6) indirekter Trommelmechanismus oder
Springfederapparat (Arius seemani). Innerhalb der Familien und Gattungen können
verschiedene Mechanismen auftreten oder fehlen. Bei den Loricariidae gibt es neben
stridulierenden Species nichtstridulierende, sogar innerhalb derselben Unterfamilie
(Ancistrinae; Schaefer 1987). Doch sind die Verwandtschaftsverhältnisse innerhalb
der Loricariidae ungeklärt. Nach Untersuchungen mit Hilfe der rRNA-Sequenzierung
sind alle Unterfamilien außer den Loricasiinae paraphyletisch (Montoya-Burgos et al.
1998).
Bei den Bagridae stridulieren Mystus gulio und M. vittatus, während A.
nemurus mit Hilfe der Kiemenspalten quietscht. Auch der Stridulationsmechanismus
kann in derselben Familie wechseln (Ictaluridae: Ameiurus, Ictalurus). Innerhalb
derselben Species treten stridulierende und nichtstridulierende Individuen auf, wobei
der Anteil der stridulierenden Exemplare unterschiedlich ist. Die einzelnen Parameter
der Geräusche wie Häufigkeit, Wiederholungsrate, Laut- und Intervalldauer, Laut-
LAUTERZEUGUNG DER WELSE (SILUROIDEI) 201
stärke oder Frequenzanteile variieren bei Familien, Arten, Angehörigen derselben
Species und sogar bei ein und demselben Individuum. Die an einer Species erhobenen
Befunde lassen sich weder für eine Familie noch für eine Gattung verallgemeinern.
Die weite Verbreitung der Lauterzeugung durch Stridulation der Brustflossen
sowie die weitgehend ähnliche strukturelle Ausbildung des Stridulationsapparates
deuten daraufhin, daß dieser Mechanismus bei manchen Familien sekundär teilweise
oder ganz verlorengegangen ist. Die Stridulationsmechanismen Abduktion und
Adduktion der Brustflossen sind sowohl gemeinsam als auch allein (nur Abduktion)
in allen größeren Verwandtschaftsgruppen zu finden (de Pinna, 1998; Tab. 3). Ent-
sprechende Untersuchungen der Basisgruppen der Welse (Diplomystidae, Cetopsidae
u.a.) liegen bisher nicht vor. Stridulationslaute treten außer bei Störungen auch beim
Fortpflanzungsverhalten mancher Species auf (Mayr, 1987; Pruzsinsky & Ladich,
1998; Kaatz & Lobel, 1999). Es gibt Hinweise, daß der Lauterzeugungsapparat bei
diesen Species sekundär abgewandelt wurde (Kaatz & Stewart, 1997).
Die Schwimmblasenmechanismen sind unterschiedlich gestaltet, doch nicht
familiencharakteristisch, wie Ladich (1997) meint. Bei den Pimelodidae besitzt
Pimelodus pictus einen Trommelmechanismus, Pimelodella gracilis nicht. Es könnte
sich dabei um einen Unterschied zwischen Unterfamilien handeln, da Pimelodus zu
den Pimelodinae, Pimelodella aber zu den Rhamdiinae (Teugels, 1996) gestellt wird.
Die letztgenannte Gruppe wird von de Pinna (1998) als Heptapterinae bezeichnet,
deren Phylogenie ungeklärt ist. Doch treten auch innerhalb derselben Unterfamilie
(Pimelodinae, Calophysus-Gruppe) Species mit und ohne Schwimmblasenmuskeln
auf (Stewart, 1986; de Pinna, 1998). Weder bei den Basisgruppen Diplomystidae,
Cetopsidae noch bei der nächsthöheren, 12 Familien umfassenden, Gruppe (Lorica-
ruidae-Callichthyidae-Aspredinidae-Trichomycteridae u.a.; de Pinna, 1998) ist bisher
ein Trommelmuskelapparat gefunden worden.
LAUTERZEUGUNG UND SCHRECKREAKTION
Die Schreckreaktion auf Schreckstoff aus der Haut von Artgenossen wurde
von Karl von Frisch (1938) an der Elritze Phoxinus phoxinus (L.) (Cyprinidae) ent-
deckt und später auch an fünf anderen Species der Cyprinidae nachgewiesen. Haut-
extrakt von 20 weiteren Species einheimischer Cyprinidae und zwei Species der
Cobitidae löste bei Elritzenschwärmen ebenfalls eine Schreckreaktion aus. Dagegen
waren die Haute von allen Fischen, die nicht zu den Cyprinidae und Cobitidae
gehören, darunter der nordamerikanische Zwergwels Ameiurus nebulosus, unwirk-
sam. Der einzige Versuch mit diesem Wels endete negativ (von Frisch, 1941a,b).
Zwei Schüler von Karl von Frisch setzten die von ihm begonnenen Versuche über die
Verbreitung der Schreckreaktion fort: F. Schutz (1956) und W. Pfeiffer (1960 bis
heute). Nach Schutz (1956) ist die Schreckreaktion unter den Cyprinidae allgemein
verbreitet, selbst wenn es sich nicht um ausgesprochene Schwarmbildner handelt.
„Die gleiche Reaktion findet sich auch bei kleineren, friedlich lebenden und
schwarmbildenden Characidae.“ Auch alle vier Versuchsgruppen von Ameiurus nebu-
losus zeigten in acht Versuchen immer ein positives Ergebnis, das Schutz (1956)
202 ANDREAS HEYD & WOLFGANG PFEIFFER
jedoch als unspezifische Reaktion „von mehr ausweichendem als schreckhaftem
Charakter“ betrachtete: „Trotz der Höhe der Konzentration der arteigenen Haut-
extrakte waren alle Reaktionen immer nur kurz und ohne nachhaltige Wirkung.“
Schutz (1956) war der Auffassung, daß eine Schreckreaktion bei den Zwergwelsen
bemerkenswert wäre, weil die jüngeren Fische in Schulen zusammenhalten. Auch die
kleinen Panzerwelse Corydoras paleatus (Callichthyidae) schwammen immer in einer
Gruppe und zeigten auf ihren Hautextrakt „leicht schreckhaftes Verhalten.“ Ihre
Reaktionsweise erzeugte bei Schutz (1956) jedoch den Eindruck, „daß es sich wie
beim Zwergwels um eine unspezifische Reaktion handelt“.
Pfeiffer (1960) zeigte in fünf Versuchen an zwei Schwärmen des indischen
Glaswelses Kryptopterus bicirrhis (Siluridae) erstmals, daß auch Siluroidei eine
Schreckreaktion besitzen, die derjenigen der Cypriniformes und Characiformes
gleichzusetzen ist. Dieses Ergebnis hat sich später an dieser und anderen Species der
Welse bestätigt (Pfeiffer, 1963a; Pfeiffer er al., 1986; Heyd & Pfeiffer, in Vor-
bereitung) und beweist, daß die Schreckreaktion bei den Ostariophysen (sensu
Sagemehl 1885) allgemein verbreitet ist. Die Entdeckung der Schreckreaktion bei den
Gonorynchitormes (Pfeiffer 1967) fügt sich in das System von Rosen & Greenwood
(1970) ein, wonach die Ostariophysen sensu Sagemehl (1885) als Otophysi mit den
Anotophysi (alias Gonorynchiformes) als Ostariophysi zusammengefaßt werden. Alle
analogen Alarm-Pheromon-Systeme von nichtostariophysen Fischen (Cottidae, Gobi-
idae, Percidae) unterscheiden sich tiefgreifend von der Schreckreaktion der Ostario-
physi (Smith, 1992). Nur die Ostariophysi (sensu Rosen & Greenwood, 1970) be-
sitzen in ihrer Epidermis Schreckstoffzellen (Pfeiffer, 1960; Pfeiffer er al., 1971),
denen der Schreckstoff entstammt (Pfeiffer, 1960, 1963a, 1967; Smith, 1973,
1976a,b).
Ob die Schreckstoffzellen noch weitere Funktionen haben, wie von Pfeiffer
(1967, 1970) für solitäre, nachtaktive Welse vermutet, ist nicht entschieden. Um die
besondere Situation der Siluroidei, was ihre Schreckreaktion anbetrifft, zu verstehen,
werfen wir zunächst einen Blick auf die anderen, wesentlich intensiver untersuchten
Ostariophysi:
(1) Innerhalb der Cypriniformes wurde die Schreckreaktion bei allen mehr als
53 bisher gepriiften Species aus fiinf Familien gefunden, mit alleiniger Ausnahme des
blinden Höhlenfisches Caecobarbus geertsi. Sie existiert bei allen tagaktiven Cyprin-
idae, auch bei bodenlebenden Species, sowie bei den Catostomidae, Cobitidae,
Gyrinocheilidae und Homalopteridae. Auch die beiden räuberisch lebenden Cyprin-
idae Leuciscus cephalus und Prychocheilus oregonense haben zumindest als Jung-
fische eine Schreckreaktion (Pfeiffer 1977).
(2) Innerhalb der Characiformes dagegen wurde die Schreckreaktion nur bei
2/3 der mehr als 50 geprüften Species gefunden. Es gibt hier Familien bzw. Unter-
familien, für welche die Schreckreaktion bisher nicht nachgewiesen werden konnte,
nämlich die Chilodontidae, Lebiasinidae, Mylinae und Serrasalminae. Während das
Fehlen der Schreckreaktion bei den räuberischen Piranhas verständlich ist, gilt dies
nicht für die ihnen nahverwandten pflanzenfressenden und friedlichen Mylinae, es sei
denn, man hält die Mylinae für Abkömmlinge der Serrasalminae. Beide, Serrasalm-
LAUTERZEUGUNG DER WELSE (SILUROIDEI) 203
inae und Mylinae, besitzen Schreckstoffzellen und Schreckstoff, doch fehlt ihnen im
ZNS die Fähigkeit, auf den Schreckstoff zu reagieren. Entsprechendes gilt für den
blinden Höhlensalmler Anoptichthys jordani (Pfeiffer, 1967, 1977).
(3) Allen untersuchten Gymnotoidei fehlen Schreckstoffzellen und Schreck-
reaktion (Pfeiffer, 1963a, 1967).
(4) Bei allen 7 histologisch geprüften Gattungen der Anotophysi aus 4 Fami-
lien wurden Schreckstoffzellen gefunden. Für die beiden im Verhaltensexperiment
getesteten Gattungen wurde die Schreckreaktion nachgewiesen. Von besonderer
Bedeutung ist die wechselseitige Wirksamkeit des Schreckstoffs zwischen Otophysi
und Anotophysi (Pfeiffer, 1967).
Von den ca. 33 Familien der Siluroidei wurden bisher 22 bezüglich ihrer
Schreckstoffzellen untersucht. Bei 20 Welsfamilien wurden sie gefunden (Tab. 3), nur
die Aspredinidae und Loricariidae führen keine (Pfeiffer, 1963c, 1977). Den Aspre-
dinidae fehlen darüber hinaus sogar die Mauthner-Axone und damit der „Mauthner-
Reflex“, mit dem jede Schreckreaktion beginnt (Göhner & Pfeiffer, 1996, Pfeiffer er
al., 1986). Im Gegensatz zu den intensiv untersuchten Cypriniformes und Characi-
formes wurden bisher nur verhältnismäßig wenige Species und Familien der Silur-
oidei bezüglich ihrer Schreckreaktion geprüft (Pfeiffer, 1977), aus verständlichen
Gründen, wie die Ergebnisse von Karl von Frisch (1938, 1941a, b) und Schutz (1956)
zeigen. Die Schreckreaktion wurde bei Angehörigen der Clariidae, Ictaluridae,
Mochokidae und Pimelodidae gesehen. Mit Sicherheit nachgewiesen, videotechnisch
aufgezeichnet und analysiert wurde sie für Kryptopterus bicirrhis (Siluridae), Pan-
gasius sutchi (Pangasiidae), Eutropiellus vanderweyeri (Schilbeidae), Corydoras
paleatus (Callichthyidae) und Plotosus lineatus (Plotosidae) (Pfeiffer, 1960, 1963a,
1977; Pfeiffer er al., 1986, Heyd & Pfeiffer, in Vorbereitung).
Die Mehrzahl der Species und Familien der Welse sind abgeflachte Boden-
fische mit hohem spezifischen Gewicht und unterständigem Maul, nachtaktiv und
kleinäugig, mit Barteln als Trägern von Tast- und Geschmacksorganen und einem
hervorragenden Hörvermögen. Sie sind meist solitär und wehrhaft durch kräftige
Stacheln der Rückenflosse und der Brustflossen, die mit Sperrmechanismen aus-
gestattet und als Stridulationsapparate prädestiniert sind. Die meisten Welse ernähren
sich von kleinen Invertebraten, einige sind Räuber, andere Algen- und Detritusfresser.
Ihre nächtliche Aktivität und ihr Bodenleben teilen sie mit den Gymnotoidei, mit
denen die Aspredinidae und Loricariidae auch das Fehlen von Schreckstoffzellen
sowie mehrere andere Welsfamilien den Besitz von Elektrorezeptoren („small pit
organs”) gemeinsam haben. Ihre Lebensweise als solitäre nachtaktive Bodenfische
unterscheidet die meisten Siluroidei und alle Gymnotoidei deutlich von den gewöhn-
lich gesellig lebenden, tagaktiven, größtenteils pelagischen Cypriniformes, Characi-
formes und Anotophysi. Die Welsarten mit der am deutlichsten ausgeprägten
Schreckreaktion sind ebenfalls tagaktive Schwarmfische oder zumindest als Jung-
fische in Gruppen lebend. Sie erinnern durch ihr Aussehen und ihre Lebensweise an
dıe große Mehrzahl der Cypriniformes und Characiformes. Corydoras ist zwar boden-
lebend, doch tagaktiv und besonders in der Jugend in Gruppen lebend. Synodontis hat
wie die Bodenfische ein unterständiges Maul, ist jedoch pelagisch und nimmt Nah-
204 ANDREAS HEYD & WOLFGANG PFEIFFER
rung von der Wasseroberfläche rückenschwimmend auf. S. nigriventris ist dement-
sprechend sogar invers pigmentiert, wie sein Name verrät. Auch junge Ictaluridae und
Pimelodidae bilden Schulen. Die pelagischen Eutropiellus, Kryptopterus, Pangasius
und Plotosus sind tagaktiv und zumindest als Jungtiere ausgesprochene Schwarm-
fische. Weil die Schreckreaktion ihre Aufgabe nur bei schwarmbildenden, tagaktiven
Fischen voll erfüllen kann, ist sie bei solitären, nachtaktiven Species nicht zu
erwarten. Der Schreckstoff dient nicht dem Sender, sondern allein den Empfängern.
Selektioniert wird nicht das Individuum, sondern die Gruppe. Nur wenn alle
Schwarmmitglieder gemeinsamer Abstammung wären, kann man von „kin“-Selektion
sprechen; meist handelt es sich um Gruppen-Selektion. Solitäre, mit kräftigen
Stacheln bewehrte Fische schützen sich selbst. Im Gegensatz zu den nachtaktiven,
meist kleinäugigen Welsen sind die tagaktiven Species großäugig, was zu der
Tatsache paßt, daß die Schreckreaktion visuell übertragen wird.
Lauterzeugung durch Stridulation ist von mindestens 14 Welsfamilien bekannt,
Trommelmechanismen von 8 Familien (Tab. 3). In 7 Familien treten beide Mecha-
nismen auf. Der elektrische Zitterwels besitzt einen Trommelapparat (Müller, 1842),
ohne zu stridulieren. Er wehrt sich mit seinen starken Entladungen, wie sonst nur
Zitterrochen und Zitteraal. Von den trommelnden und stridulierenden Familien be-
sitzen die Mochokidae und Pimelodidae die Schreckreaktion, von den nur stridu-
lierenden die Ictaluridae und Callichthyidae. Während den stridulierenden Aspredin-
idae und Loricariidae Schreckstoffzellen und Schreckreaktion fehlen, sind von den
pelagischen Welsen mit besonders gut ausgebildeter Schreckreaktion nur vereinzelt
Stridulationsmechanismen bekannt. Schreckreaktion und Lauterzeugung schließen
einander zwar nicht aus, aber es ist auffallend, daß sie entsprechend der Lebensweise
der Fische verbreitet sind. Viele nachtaktive, bodenlebende, wehrhafte, solitäre Welse
können durch Stridulation Abwehrlaute erzeugen. Die tagaktiven, schwarmbildenden
Welse zeigen eine Schreckreaktion. Doch ist es gegenwärtig wenig ratsam, einen nur
an wenigen Species erhobenen Befund für eine ganze Familie zu verallgemeinern.
Aufgrund unvollständiger Untersuchung ist unser bisheriges Bild lückenhaft. Deshalb
sollten Schlüsse über den Zusammenhang zwischen dem Auftreten von Lauterzeu-
gungsmechanismen einerseits und dem Vorkommen der Schreckreaktion andererseits
nur mit Vorsicht gezogen werden.
DANKSAGUNG
Wir danken Dr. R. Britz und Dr. H.-J. Franke für Hilfe in taxonomischen
Fragen sowie Prof. Dr. R. Apfelbach, Prof. Dr. E. Müller, Dr. R. Triebskorn,
H. Casper und I. Kaipf für technische Unterstützung.
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REVUE SUISSE DE ZOOLOGIE, 107 (1): 213-232; mars 2000
Molecular and morphological evidence for the presence of a
new Buthid taxon (Scorpiones: Buthidae) on the Island of Cyprus
Benjamin GANTENBEIN*!, Christian KROPF?, Carlo Rodolfo LARGIADER!
& Adolf SCHOLL!
| Division of Population Biology, Institute of Zoology, University of Berne,
Baltzerstrasse 3, CH-3012, Berne, Switzerland.
e-mail: Gantenbein @zoo.unibe.ch
2 Natural History Museum Berne, Department of Invertebrates,
Bernastrasse 15, CH-3005 Berne, Switzerland.
e-mail: christian.kropf@nmbe.unibe.ch
Molecular and morphological evidence for the presence of a new
Buthid taxon (Scorpiones: Buthidae) on the Island of Cyprus. - Allo-
zyme data (16 loci) from Mesobuthus gibbosus populations in the eastern
Mediterranean region show that the Cyprus population is highly distinct,
although morphological differentiation is rather weak. This provides
evidence for a ‘hidden’ taxon on Cyprus. This island population 1s described
as a new species.
Key-words: Allozymes - Scorpiones - Mesobuthus — new species - Cyprus.
INTRODUCTION
Comparison of molecular data from insular and continental populations of a
species or of closely related species may provide new insights into the process of
speciation with respect to time (Estoup er al., 1996; Baldwin & Sanderson, 1998; Beerli
et al., 1996; Gillespie et al., 1998; Hollocher, 1998; Widmer er al., 1998). Island
populations are of particular interest for the analysis of colonisation processes (Vachon
& Abe, 1988) or for studying founder effect speciation (Templeton, 1980, 1981; Carson
& Templeton, 1984; Grant, 1998). Mesobuthus gibbosus (Brullé, 1832) (Scorpiones:
Buthidae) is widely distributed in the eastern Mediterranean region (Werner, 1928,
1937, 1938; Vachon, 1947a, 1947b, 1948, 1966; Tolunay, 1959; Gruber, 1963, 1966;
Kinzelbach, 1975, 1982, 1984, 1985; Vachon & Kinzelbach, 1987; Kritscher, 1993;
Crucitti, 1993). Its geographical range includes the mainland (south Balkan, Pelo-
ponnesus, Anatolia) and eastern Mediterranean islands (e. g. Cyclades, Sporades, Crete
“to whom correspondence should be addressed
Manuscript accepted 05.10.1999
214 B. GANTENBEIN, C. KROPF, C. R. LARGIADER & A. SCHOLL
and Cyprus). The geographic variation within scorpion species was traditionally exami-
ned by using morphological characters like patterns of trichobothria (=‘tricho-
bothriotaxy’) (Vachon, 1974, 1975, 1976; Fet, 1986; Fet & Rechkin, 1989) or variation
in the number of pectinal teeth (Kinzelbach, 1975; Michalis & Kattoulas, 1981;
Michalis & Dolkeras, 1989). The taxonomic status of different populations of M.
gibbosus has remained unclear: Kinzelbach (1975) distinguished two subspecies, M. g.
gibbosus (Brullé, 1832) (south of the Balkan Peninsula, Northern Sporades, Cyclades)
and M. g. anatolicus (Schenkel, 1947) (Crete, Anatolia, Cyprus). Kritscher (1993)
doubted the validity of M. g. anatolicus and rejected it.
The widely accepted theory of the salinity crisis (Hsü, 1972; Hsü et al., 1977)
states that the Mediterranean sea dried out 5.6 Myrs ago and, consequently, the
colonisation of islands via landbridges was possible during a period of 100°000 yrs.
Since the refilling of the basin (5.2 Myrs ago) the populations became geographically
isolated. Since then, a considerable degree of differentiation between mainland and
island populations is expected to have taken place. Alternatively, other genetic
population structures than expected are likely if scorpions were introduced by man.
Because of several introductions of euscorpiids (Stockwell 1992) caused by man (e. g.
Benton, 1991; Goyffon, 1992; Toscano-Gadea, 1998), island populations might not be
as isolated as expected.
To clarify the status of some island populations and of the Cyprus population in
particular, we carried out a comparative genetic analysis at 16 allozyme loci.
Populations from the Peloponnesus, from the south of the Balkans, from Crete, Rhodes,
Anatolia and from Cyprus were included. We describe here new morphological
characters, which distinguish Mesobuthus from Cyprus from the other populations of
Mesobuthus gibbosus examined.
MATERIAL & METHODS
Specimens analysed. Samples were collected at two sites on the Peloponnesus,
GR (Vigla [Arta]; Mathia [Messinia]), two sites on Crete, GR (Vai; Zakros [both in
Lassithi]), two sites on Rhodes, GR (Petaloudes: Kolympia), two sites in central
Anatolia, TR (Avanos; Hacibectas [both Cappadocia]), one site in southern Anatolia,
TR (Selale [Pamphylia]) and two sites on northern Cyprus (Tepebasi; Kantara [both
Turkish part]). The scorpions were transported alive to the laboratory, killed by deep-
freezing and stored at -80°C prior to electrophoresis. For the morphological exami-
nation, the specimens were later transferred into 75% ethanol. Androctonus maure-
tanicus (Pocock, 1902) from Agadir (MA) and Androctonus crassicauda (Olivier,
1807) from Urfa (TR) were used as outgroup species for phylogenetic analyses. The
sampling sites are shown in Fig. 1.
Allozyme analysis. Horizontal starch gel electrophoresis of allozymes was
carried out according to the protocols described in Gantenbein er al. (1998) and Murphy
et al. (1996). We scored 16 loci on three buffer systems: N-(3-Aminopropyl)-morpho-
line-citrate (AC, pH 6.2, modified from Clayton & Tretiak, 1972), Tris-citrate (TC, pH
7.3, Ayala et al., 1972) and Tris-borate-EDTA (TBE, pH 9.3, modified from Ayala er
A NEW BUTHID TAXON ON THE ISLAND OF CYPRUS
(N°)
Nn
Zo &
The BALKANS
=
\
ne
SS) 5
4
FIG. |
Sampling sites of the analysed Mesobuthus populations: 1: Vigla, GR, 2: Mathia, Gr, 3: Vai, GR,
4: Zakros, GR, 5: Kolympia, GR, 6: Petaloudes, GR, 7: Selale, TR (3 sites), 8: Hacibectas, TR, 9:
Avanos, TR, 10: Kantara, TR, 11: Tepebasi, TR.
al., 1972). The loci scored were: ALPDH (Alanopine dehydrogenase; EC 1.5.1.17),
ARK (arginine kinase; EC 2.7.3.3), AAT-1 and AAT-2 (aspartate aminotransferase; EC
2.6.1.1), DDH (dihydrolipoamide oxidase; EC 1.8.1.4), PGI (GPI) (glucose-6-phos-
phate isomerase; EC 5.3.1.9), GTDH (glutamate dehydrogenase; EC 1.4.1.2), IDH-1
and IDH-2 (isocitrate dehydrogenase; EC 1.1.1.42), MDH-I and MDH-2 (malate
dehydrogenase; EC 1.1.1.37), MPI (mannose-6-phophate isomerase; EC 5.3.1.8), PGM
(phosphoglucomutase; EC 5.4.2.2), 6-PGD (6-phosphogluconate dehydrogenase; EC
1.1.1.44), PK (pyruvate kinase; EC 2.7.1.40), and SOD (superoxide dismutase; EC
1.15.1.1). We refer to the observed electromorphs as alleles which are identified by
their electrophoretic mobility relative to the most common mobility in the Euscorpius
flavicaudis (de Geer, 1778) population from Lauris, France (assigned mobility=100) as
described in Gantenbein er al. (1998). To assess the genetic variability within each
population, the mean number of alleles per locus, the percentage of polymorphic loci
and the mean heterozygosity were calculated by the direct count method and by Nei’s
(1978) unbiased estimate. Nei’s genetic distance (1972) was calculated from pairwise
216 B. GANTENBEIN, C. KROPF, C. R. LARGIADER & A. SCHOLL
comparisons of populations using the program GENDIST from the PHYLIP 3.5
package (Felsenstein, 1995). Nei’s distance is expected to rise linearly with time since
the complete separation of gene pools, if sufficiently large numbers of loci are
measured (Felsenstein 1984; Nei 1987). Using Nei’s pairwise distances as an input
matrix, an additive tree was created by the neighbour-joining algorithm (NJ) (Saitou &
Nei, 1987). NJ is assumed to be a good heuristic approach for estimating the minimum
evolution tree (Page & Holmes, 1998) and allows for different rates among lineages in
contrast to the UPGMA clustering algorithm. Alternatively, an unrooted maximum
likelihood tree was calculated using the computer program CONTML. It estimates
phylogenies by the restricted maximum likelihood (REML) method, based on the
Brownian motion model (Cavalli-Sforza & Edwards, 1967). The REML algorithm was
formerly described in Felsenstein (1973, 1981) and uses less parameters than the full
ML analysis and is, therefore, assumed to be more consistent. Additionally, the
program calculates branch lengths and rough confidence intervals for the branches.
Bootstrap values were obtained based on 1000 pseudo-replicates of allele frequencies
using the SEQBOOT routine in PHYLIP.
RESULTS
Genetic variability. In general, the observed genetic variability was rather low
within the Mesobuthus samples. The mean observed heterozygosity was 0.07 + 0.05,
one to six out of 16 loci were polymorphic at the 0.95 criterion. The mean number of
alleles ranged from 1.0 to 1.5 per locus. All analysed populations were fixed at many
loci (Table 1). However, this variation was not evenly distributed among Mesobuthus
samples. At five loci, more than two electromorphs were detected, whereas all
populations were fixed for the same allele at three loci (Idh-2, Mdh-1, Pk). The Cyprus
samples were fixed for private alleles (alleles that were not found in any other
population) or showed polymorphisms with private alleles at six loci (Aat-1, Aat-2,
Alpdh, Ldh-2, Mdh-2, 6-Pgd) in addition to minor differences in allele frequencies at
the locus Mpi (Table 1). On the other hand, if compared to the mainland populations,
the island populations from Crete and Rhodes showed private alleles at one locus only
(samples Zakros and Vai at Aat-1 and samples Kolympia and Petaloudes at 6-Pgd,
respectively). The island samples and the mainland samples differed mainly in the
allele frequencies.
The outgroup comparison with A. mauretanicus and A. crassicauda, respecti-
vely, revealed alleles at eight loci that were not found in Mesobuthus, whereas, with
respect to allele frequencies, the two Androctonus species differed from each other only
at two loci.
TABLE |
Allele frequencies and sample sizes of eleven Mesobuthus populations. Androctonus maure-
tanicus (MA) and A. crassicauda (TR) were used as outgroup species. Alleles were labelled as
described in the material & methods section. Measures of genetic variability for each population
are given at the bottom of the table. Private alleles of the Cyprus samples and the heterozygosity
Genus/species
Region
Site
Sample size
Locus Allele
Aat-1 111
Aat-2 117
Alpdh 105
Ark 102
Ddh 99
Gtdh 100
Idh-1 104
Idh-2 93
Mdh-1 105
Mdh-2 92
Mpi 160
6-Pgd 112
Pgi (GPI) 95
87
Pgm 88
81
Pk 100
98
Sod 110
109
103
Mean no. of
alleles per locus
Percentage of
polymorhphic loci
Heterozygosity
(observed)
Heterozygosity
(expected)
Greek mainland
A NEW BUTHID TAXON ON THE ISLAND OF CYPRUS
M. gibbosus
Crete
Vigla Mathia Zakros Vai
(2)
1.00
1.00
0.03
1.00
1.00
1.00
1.00
1.00
1.00
1.00
1.00
0.25
0.25
0.50
1.00
0.25
0.75
1.00
1.00
0.06
(0.0)
0.08
(0.1)
(15)
1.00
0.97
1.00
1.00
1.00
1.00
1.00
1.00
1.00
1.00
1.00
1.00
(5)
1.00
1.00
1.00
1.00
1.00
1.00
1.00
1.00
1.00
1.00
0.50
0.50
1.00
1.00
1.00
1.00
1.00
1.1
(0.0)
6.3
0.01
(0.0)
0.04
(0.0)
(2)
1.00
1.00
1.00
1.00
1.00
1.00
1.00
1.00
1.00
1.00
1.00
1.00
Rhodes
bia
(6)
1.00
1.00
1.00
0.67
0.33
1.00
0.67
0.33
1.00
1.00
1.00
1.00
1.00
0.08
0.92
Kolym- Peta-
loudes
(6)
1.00
1.00
0.09
1.00
22
oo —
wn
1.00
0.25
0.75
0.08
0.92
1.00
1.00
1.00
1.00
S. Ana-
tolia
Selale
(16)
1.00
0.91
1.00
1.00
1.00
1.00
0.81
0.19
1.00
1.00
1.00
0.97
0.03
1.00
C. Ana-
tolia
Avanos Haci-
bectas
(5) (3)
1.00 1.00
1.00 1.00
1.00 1.00
1.00 1.00
1.00 1.00
1.00 1.00
0.90 1.00
0.10
1.00 1.00
1.00 1.00
1.00 1.00
0.90 1.00
0.10
1.00 1.00
1.00 1.00
1.00 1.00
1.00 1.00
1.00 1.00
1.1 1.0
(0.1) (0.0)
12.5 0.0
0.03 0.0
(0.0) (0.0)
0.03 0.0
(0.0) (0.0)
M. cyprius
Cyprus
Tepe- Kan-
basi tara
(20) (4)
0.30
0.70 1.00
0.35 0.63
0.13
0.65 0.24
1.00 1.00
1.00 1.00
0.05
0.95 1.00
1.00 1.00
0:977721:00
0.03
1.00 1.00
1.00 1.00
1.00 1.00
0.62 1.00
0.38
0.03
0.12
0.94 0.88
0.03
0.08
0.92 1.00
1.00 1.00
1.00 1.00
1.00 1.00
1:5 12
(0.2) (0.1)
Is NAS
0.08 0.06
(0.0) (0.0)
0.11 0.05
(0.1) (0.0)
Aourir
(14)
1.00
1.00
1.00
1.00
1.00
1.00
1.00
1.00
1.00
0.04
0.96
1.00
1.00
0.03
(0.0)
0.04
(0.0)
217
Androctonus
Morocco Turkey
Urfa
(6)
0.08
0.75
0.17
1.00
1.00
1.00
1.00
1.00
1.00
1.00
1.00
0.67
0.33
1.00
1.00
0.09
(0.1)
0.09
(0.1)
218 B. GANTENBEIN, C. KROPF, C. R. LARGIADER & A. SCHOLL
TABLE 2
Calculated pairwise distance matrix (Nei’s D 1972) for all 13 samples of Mesobuthus. In bold are
the genetic distance values of pairwise comparisons that involved one of the two Cyprus samples.
Region Greek mainland Crete Rhodes Anatolia Cyprus Outgroup
Population Vigla Mathia Zakros Vai Kolympia Peta- Selale Avanos Haci- Tepe- Kantara Aourir
loudes bectas basi
Vigla
Mathia 0.03
Zakros 0.42 0.41
Vai 0.44 0.47 0.02
Kolympia 0.13 0.10 0.21 0.26
Petaloudes 0.05 0.03 0.35 0.42 0.05
Selale 0.45 0.39 0.16 0.21 0.19 0.33
Avanos 0.34 0.29 0.23 0.28 0.11 0.24 0.07
Hacibectas 0.34 0.29 0.23 0.29 0.10 0.23 0.07 0.00
Tepebasi 0.31 0.28 0.78 0.84 0.44 0.33 0.87 0.70 0.69
Kantara 0.39 0.33 0.82 0.92 0.50 0.39 0.95 0.78 0.77 0.03
Aourir 1.40 [ESI 1.13 IIS 1.09 1.32 0.95 0.80 0.81 1.32 135
Urfa 1.49 1.55 1.26 1.28 1.11 1.30 1.04 0.88 0.90 1.44. 1.46 0.14
Genetic differentiation and phylogenetic analyses. The calculated Nei’s dis-
tances from pairwise comparisons of populations within Mesobuthus ranged from 0.00
to 0.95 (Table 2). The distance values between the two outgroup species and Meso-
buthus were at least 0.80 and ranged up to 1.55. If the two samples from Cyprus are
compared with the other Mesobuthus samples, it is obvious that the Cyprus populations
are considerably distinct from all others, 1. e. Vigla, Mathia, Kolympia, Petaloudes,
Zakros, Vai (all GR), Selale, Avanos, Hacibectas (all TR), by rather high distance
values (0.31 - 0.95).
The topologies of the NJ tree and of the maximum likelihood (ML) tree (-In
likelihood 544.123; 4909 trees examined) are generally congruent and differ from each
other only in the position of the Cretean clade (Figs 2-3). This clade clusters separately
in the NJ tree (Fig. 2) and groups with the sample of Selale (south Anatolia) in the ML
tree (Fig. 3). However, both approaches revealed four main clades within Mesobuthus
(Figs 2-3). The first clade is composed of the samples from the mainland of Greece
(Vigla and Mathia) and island Rhodes (Petaloudes and Kolympia) and is found at a
relatively low distance value. The two samples from Crete are found in a second
independent clade, and the populations from Anatolia as a third clade. However, the
Cyprus clade remains clearly separated from all other clades by a rather high tree length
in both trees, and is confirmed as an offshot from the mainland. The bootstrap analysis
supports four nodes of the trees with very high values (>90%), these are observed for
the clades grouping the Cyprus samples (Tepebasi and Kantara), the central Anatolian
samples (Avanos and Hacibectas), the Crete samples (Zakros and Vai), and the
Androctonus species, respectively. The node between the Peloponnesian clade and the
Cyprus clade is well supported (60% and 77%, respectively). Androctonus is confirmed
as outgroup by a bootstrap value of 99%. However, the other nodes are weakly
supported. The weakest support is found for the clade that contains the Cretean and
Anatolian samples. Noteworthy, the outgroup species A. mauretanicus and A. crassi-
cauda are separated by about the same genetic distance that is found among the samples
A NEW BUTHID TAXON ON THE ISLAND OF CYPRUS 219
of the Peloponnesus / Rhodes clade and the Anatolian clade, although the outgroup
samples are treated as different species and are geographically separated by a large
distance (approximatively 3000 km).
Avanos
Mesobuthus Anatolia
Hacibectas
Selale
Zakros
Vai Crete
Vigla
Mathia
Peloponnesus
Cyprus |
Kantara
96! Tepebasi
Petaloudes
Kolympia Rhodes
Androctonus Urfa ——
0.1 0
91 Aourir Nei's D
FIG. 2
NJ tree based on Nei’s distance relating island and mainland populations of Mesobuthus inferred
from 16 allozyme loci. A. crassicauda from Urfa (TR) and A. mauretanicus from Aourir (MA)
were used as outgroups.
TAXONOMY
Mesobuthus cyprius Gantenbein & Kropf, sp. n. Figs 4-28; Tab. |
Type material: Holotype: 1 male, Tepebasi, Northern Cyprus, 26. V. 1998,
Natural History Museum Berne, Switzerland (NMBE). Paratypes (collected at the type locality):
I male, 20. V. 1998, 1 female, 26. V. 1998, NMBE. 1 male, 20. V. 1998, 1 female, 26. V. 1998,
Natural History Museum Basel, Switzerland; all specimens leg. A. Scholl (Berne, CH).
Other material examined: Morphology: Mesobuthus cyprius sp. n.: 6 males,
11 females from Tepebasi, I male, 1 female from Kantara, Cyprus. For a comparison, also
specimens of Mesobuthus gibbosus from Rhodes, GR (31 specimens), Peloponnesus, GR (7),
Crete, GR (20), Euboea, GR (1), Selale, TR (12) and central Anatolia, TR (3) were investigated.
Allozyme study: Mesobuthus cyprius sp. n., Mesobuthus gibbosus,
Androctonus mauretanicus, Androctonus crassicauda: See Table 1.
Diagnosis: Mesobuthus cyprius Sp. n. can be distinguished unambiguously
only by the shape of the basal lobes of the hemispermatophores. These are slender and
acutely pointed teeth in Mesobuthus cyprius sp. n. (Figs 8-9), while in all populations of
M. gibbosus examined they form scales with more or less distinct blunt tips (Figs 29-
220 B. GANTENBEIN, C. KROPF, C. R. LARGIADER & A. SCHOLL
Vai
99! Zakros Crete
Selale
Mesobuthus Avanos Anatolia
Hacibectas
4
> Vigla
Peloponnesos
Mathia
Kantara
99 | Tepebasi Cyprus
Petaloudes
Kolympia Rhodes
Androctonus Urfa
89 Aourir
freni
0.1 0
REML distance
Fic. 3
Rooted maximum likelihood (ML) tree of island and mainland populations of Mesobuthus.
33: Vachon, 1948: fig. 1). Moreover, the hemispermatophore of the new species is
considerably smaller than in M. gibbosus.
Description: Measurements (in mm): No apparent size dimorphism,
although females longer, and males with a longer metasoma. Total length (measured
dorsally from anterior margin of carapace to tip of stinger, with telson in horizontal
position): Males 45-55, females 45-60. Carapace length (measured dorsally along
midline): males 5.0-5.7, females 5.0-6.0. Carapace width (maximum distance between
posterio-lateral edges): males 5.5-6.0, females 5.5-6.6. Metasoma length: males 28-37,
females 27-35 (measured dorsally from anterior margin of first segment to tip of
stinger).
Carapace (Figs 4, 5): trapezoid-shaped, colour pattern more or less distinct,
especially lateral pigmented areas hardly visible in some specimens. Granulation: ante-
rior median, central median and central lateral carinae distinct, granulae in remaining
areas mostly weak. Anterior margin with transverse row of bristles.
Mesosoma (Figs 5, 6): Tergites I-VI with a pattern of pigmented longitudinal
stripes, i.e. one median stripe and two pairs of lateral stripes (one paramedian, one ectal
pair). These stripes partly broken, especially on tergite 6, colour varying from dark grey
to medium brown (only alcohol preserved material examined, colour pattern more
distinct in juveniles than in adults, in some adults hardly visible). Interspace between
median and paramedian stripe less than 1-1,5 times as wide as paramedian stripe at
posterior margin of tergites IV-VI. Tergite VII with more or less pronounced median
A NEW BUTHID TAXON ON THE ISLAND OF CYPRUS 22]
3
+ 59)
7
Fic. 4-6
Mesobuthus cyprius sp. n. — 4. Female carapace, dorsal view. — 5. Male carapace and mesosoma,
dorsal view. — 6. Sternum and genital opercula, ventral view.
B. GANTENBEIN, C. KROPF, C. R. LARGIADER & A. SCHOLL
(O)
[59]
(N°)
keel bearing granula. Tergites only sporadically with bristles or without. Sternites
smooth, at least some of them with white guanin spots: these spots sometimes distri-
buted all over the ventral and occasionally also the dorsal surface. Sternum subtrian-
gular, with deep median depression flanked by a pair of bristles (Fig. 6). Genital
opercula with a few scattered bristles (Fig. 6). Pectines with 20-23 (females) or 26-30
(males) teeth.
Hemispermatophore (Figs 7-9): Long, slender, of the flagelliform type. Four
lobes at truncal flexure: internal lobe biggest, with a small distal hook, close to the
external lobe, partly embracing it. Basal lobe developed as a slender and acutely
pointed tooth.
Metasoma (Figs 10-12): Segments I-IV with 10 longitudinal keels formed by
rows of tubercles; one additional pair of keels mediodorsally being indistinct and hardly
visible (not illustrated). Fourth segment 1.79 — 2.44 times longer than high in males,
1.54 — 1.58 times in females (length measured dorsally, height measured as distance
between dorsal and ventral keel). Segment V granulate ventrally, with 3-4 blunt teeth
laterally at distal edge. Telson with dark tip of stinger, granulate on ventral side, with
several short bristles, one ventral pair of them rising from subaculeate tubercle.
Chelicera (Figs 13-14): Tibia (nomenclature following Hjelle 1990) with two
large bristles dorsaliy near distal-prolateral edge, three shorter translucent hairs (several
more of them on movable finger) and a brush of fine hairs prolaterally. Fixed finger
with two ventral and four dorsal teeth; medial and basal dorsal tooth fused to form a
bicusp. Movable finger with five dorsal teeth, the two basal ones being the smallest;
ventrally with three teeth, i.e. one distal and two medial ones.
Pedipalp (Figs 15-23): Trichobothrial pattern constant, corresponding to type A
of Vachon (1974). Femur a little shorter than patella, with 11 trichobothria, d2
(nomenclature of trichobothria following Sissom, 1990) short and translucent (Fig. 17).
Patella with two distinct tooth-like tubercles ventro-proximally and with 13 tricho-
bothria, d2 short and translucent (Figs 18-19). Chela slender, males with a weak lobe on
movable finger (not illustrated). 15 trichobothria present; esb, Esb and Eb3 short and
translucent (Figs 20-21). Teeth on inner side of fingers arranged in a characteristic
pattern, interspersed with short, stiff bristles, i.e. groups of two retrolateral and one
prolateral bristle and several of them at the tip (Figs 22-23).
Legs (Figs 24-28): increasing in length from leg | - 4. All legs with a pair of
ventral pedal spurs distally on basitarsus. Proventral spur carrying a hair and a
translucent tooth originating proximally on it (Fig. 28). Retroventral spur with a small
tooth close to spur (Fig. 27). Legs 3 and 4 with a tibial spur distally (Fig. 26). Dorsal
distitarsus with a conspicuous distal hair on an elevated base (Fig. 28). Apotele with
three claws (Fig. 27).
Remarks: Mesobuthus cyprius sp. n. differs from most populations of M.
gibbosus by its pigmentation pattern on the mesosomal tergites, particularly by the
width of the pale interspace between the median and the paramedian dark stripe which
is less than 1-1,5 times the width of the latter at the posterior margin of tergites IV-VI.
In most populations of M. gibbosus, the width of this interspace is at least more than 2
times the width of the paramedian stripe which may even be absent. However, this
i)
(NO)
(SS)
A NEW BUTHID TAXON ON THE ISLAND OF CYPRUS
0,5 mm
> GE
Se
Se > A
Fics 7-12
Mesobuthus cyprius sp. n. — 7. Left hemispermatophore, total view; surrounding tissue of paraxial
organ removed, except in distal part of flagellum. — 8, 9. Lobes at truncal flexure. — 10.
Metasoma segments 4, 5, and telson of male, dorsal view. — 11. Ditto, lateral view. — 12. Ditto,
ventral view.
224 B. GANTENBEIN, C. KROPF, C. R. LARGIADER & A. SCHOLL
WA
ZI I (1 A
i LA
Fic. 13-16
Mesobuthus cyprius sp. n., male — 13. Right chelicera, ventral view; prolateral hair brush
simplified. — 14. Ditto, dorsal view. — 15. Right pedipalp, ventral view. — 16. Ditto, dorsal view.
A NEW BUTHID TAXON ON THE ISLAND OF CYPRUS 225
DN
ù SN EN alte ele
Le.) a, NEE BZ
oo, 20800 5608020 es
Cesa =
z Ge — Cai 0,5 mm
22
Fics 17-23
Mesobuthus cyprius sp. n., male — 17. Right palpal femur, trichobothrial pattern, dorsal view. —
18. Right palpal patella, trichobothrial pattern, dorsal view. — 19. Ditto, retrolateral view; arrow
and asterisk in Figs 18, 19 indicate the same trichobothria. — 20. Right chela, trichobothrial
pattern, proventral view. — 21. Ditto, retrodorsal view. — 22. Distal part of fixed finger of right
chela, inner side. — 23. Tip of movable finger of right chela, inner side.
226 B. GANTENBEIN, C. KROPF, C. R. LARGIADER & A. SCHOLL
Fics 24-28
Mesobuthus cyprius sp. n., male legs. — 24. Left leg I, prolateral view. — 25. Ditto, retrolateral
view. — 26. Left leg IV, prolateral view. — 27. Left tarsus I, ventral view; hairs omitted. — 28.
Ditto, dorsal view.
A NEW BUTHID TAXON ON THE ISLAND OF CYPRUS DDT
2
2.
Bar 30. Le
0,5 mm
IEZZO
BO as 33
Fics 29-33
Mesobuthus gibbosus, lobes at truncal flexure of left hemispermatophore in specimens from
different regions; same magnification as in Figs 8-9. — 29. Euboea (GR). — 30. Rhodes, Kolympia
(GR). — 31. Rhodes, Monolithos (GR). — 32, 33. Selale (TR).
228 B. GANTENBEIN, C. KROPF, C. R. LARGIADER & A. SCHOLL
distinction is not reliable. On the one hand the mesosomal pattern is hardly visible in
some animals, on the other hand specimens of M. gibbosus from the Peloponnesian
peninsula (Mathia) and from Crete can also show this pigmentation pattern. The
Peloponnesian individuals of M. gibbosus differ by a characteristic row of bristles on
the mesosomal tergites, which cannot be found in Mesobuthus cyprius sp. n. There,
only some scattered bristles, if any at all, do occur. Some of the Cretean specimens of
M. gibbosus examined cannot be distinguished from M. cyprius sp. n. by means of
external morphology, because they possess a similar pigmentation pattern and lack
mesosomal bristle rows.
DISCUSSION
The allozyme data generally revealed a low genetic variability (Table 1), as
already observed in earlier studies on scorpions of the family Euscorpiidae (Stockwell
1992; Gantenbein ef al., 1998, 1999). One exception is the population of M. gibbosus
from the island of Rhodes which shows a higher level of heterozygosity as compared to
the other Mesobuthus populations. This is mainly due to three gene loci (Ark, Gtdh,
Sod) (Table 1). Specimens of M. gibbosus from Rhodes that are heterozygous at these
three loci possess one allele, which is elsewhere only found in the samples from the
Peloponnesus, and another allele, which is common in the Cretean and Anatolian
populations. We speculate that this pattern indicates hybridization between western
Aegean populations and eastern Aegean populations on the island of Rhodes. This
hybridization may have resulted from secondary contact between the authochtonous
Rhodes population and an introduced population of Greek origin.
The allozyme gene frequencies indicate that there are different population
groups within Mesobuthus. In the NJ and ML tree, the Cyprus clade is separated from
the other clades by a long branch (Figs 2-3). Both approaches clearly underline that the
Cyprus scorpions have diverged considerably. Since the Biological Species Concept
(BSC) (Mayr 1942) can hardly be applied to allopatric populations, it remains unsolved
if the Cyprus population represents a separate species in this sense. Consequently, there
are no objective arguments for the taxonomic rank of this population. However, our
expectation that this island population remained isolated from the mainland populations
since the colonisation 5.2 Myrs ago is supported by a high genetic divergence of the
Cyprus clade and by the noninterchangeable morphology of the hemispermatophores.
Taking these two arguments into account we, therefore, designate to the Cyprus
population the rank of a species.
It is remarkable that M. cyprius sp. n. appears genetically distinct and highly
differentiated, but can hardly be characterised by means of classical morphology. Such
‘hidden’ taxa are well known from other animal groups (e. g. mosquitos, polychaetes,
myriapodes) (Narang ef al., 1989), especially on islands. The only characters that can
be used for a morphological distinction are found in the hemispermatophores, in the
mesosomal pigmentation pattern and in a lack of mesosomal bristle rows. The last
character is a negative trait and presumably represents a plesiomorphic state as
compared to most populations of M. gibbosus. The bristle row was found in specimens
A NEW BUTHID TAXON ON THE ISLAND OF CYPRUS 229
of M. gibbosus from the Peloponnesus (Mathia) and from Rhodes (Petaloudes). The
pigmentation pattern seems to be quite useful, because we found only limited variation
there (but compare paragraph ‘remarks’ above). The taxonomic value of the pigmen-
tation pattern was underlined by Lourenco (1983), who claimed that species with poly-
morphic colouration or pigmentation are an exception among buthids. As a conclusion,
only a single reliable distinctive morphological character of M. cyprius sp. n. remains
discriminant, 1.e. the shape of the basal lobes of the hemispermatophores.
Based on a single male, Schenkel (1947) described a new variety of M. gibbosus
from central Anatolia (collected along the road between Sivas and Amasya, Turkey).
Kinzelbach (1975) elevated Schenkel’s variety to subspecies rank. He distinguished
M. g. gibbosus (Brullé, 1832) in the south of the Balkan Peninsula, Northern Sporades,
Cyclades, from M. g. anatolicus (Schenkel 1947) living in Crete, Anatolia, and Cyprus,
using differences in the number of pectinal teeth as the main character. Furthermore, he
argued that these two subspecies were separated since the Miocene (12-25 Myrs BP).
Kritscher (1993) refuted Kinzelbach’s arguments for a separate subspecies anatolicus
and rejected this taxon. The allozyme data now partially support the existence of two
subspecies in Kinzelbach’s sense. The Greek mainland populations are genetically
slightly distinct from the Crete-Anatolian population group (Figs 2, 3). However, the
Cyprus population (= Mesobuthus cyprius sp. n.) is genetically considerably more
differentiated from all others, as discussed above. This fact contradicts the existence of
M. g. anatolicus on Cyprus as proposed by Kinzelbach.
ACKNOWLEDGEMENTS
Rica and Andreas Quensel, Ahmet Ylzaz and Keco Kutlay, Girne (Northern
Cyprus) provided logistics for field work and collecting in Northern Cyprus and
Turkey. Iasmi Stathi, Matt Braunwalder and Peter Schwendinger (Muséum d’histoire
naturelle, Geneve, CH) contributed specimens from Crete. Field trips were financially
supported by the Dr. Karl Bretscher-Foundation Berne. Beatrice Liischer and Lilian
Beer assisted in the allozyme analysis. The comments of Victor Fet and two anonymous
reviewers greatly improved the final version of the manuscript.
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REVUE SUISSE DE ZOOLOGIE
Tome 107 — Fascicule |
JUVARA-BALS, Ilinca & Vojciech WITALINSKI. Description of five new
species of Holoparasitus s. str. with redescription of H. apenninorum
(Berlese, 1906) and H. cultriger (Berlese, 1906) from Italy and Spain
(AcangGamasida Para side NE PER RO
SiLva, Carla Maria Menegola da & Beatriz MOTHES. Three new species of
Geodia Lamarck, 1815 (Porifera, Demospongiae) from the bathyal
depths off Brazilian coast, Southwestern Atlantic.................
MAHUNKA, Sandor & Luise MAHUNKA-Papp. Oribatids from Switzerland III
(Acari: Oribatida: Oppiidae | and Quadroppiidae). (Acarologica Ge-
GV CHEST ODN CNN a I Us
GIL DE PERTIERRA, Alicia A. & Alain de CHAMBRIER. Rudolphiella szidati
sp. n. (Proteocephalidea: Monticelliidae, Rudolphiellinae) parasite of
Luciopimelodus pati (Valenciennes, 1840) (Pisces: Pimelodidae) from
Argentina with new observations on Rudolphiella lobosa (Riggenbach,
1006): a N en CE E ce Gene
BENJAMIN, Suresh P. & Rudy JOCQUÉ. Two new species of the genus
Sujjasia trom Sr Lanka (Araneae: Zodariidae). . dr Re
LANG, Claude. Diversité du zoobenthos dans 47 rivières du canton de Vaud:
temaance yO SIERO Re N RE ES
VIT, Stanislav. Contribution a la connaissance de la famille Eucinetidae
(COEO OS) ARS EE eno DR PERU ev Wee Cia
BÄNZIGER, Ruth. Spatio-temporal distribution of size classes and larval
instars of aquatic insects (Ephemeroptera, Trichoptera and Leopi-
doptera) in a Potamogeton pectinatus L. bed (Lake Geneva, Switzer-
Fann) reenact ee Wee el NG NER SINE NT Vogt RELA,
BESUCHET, Claude & Stanislav VIT. Les Nanophthalmus Motschulsky
eEurope (Coleoptera Scydmacnidac))a ri 220, nen,
HEYD, Andreas & Wolfgang PFEIFFER. Uber die Lauterzeugung der Welse
(Siluroidei, Ostariophysi, Teleostei) und ihren Zusammenhang mit der
Biyloseniesund der Schreckrealsti ona 2... nun. on:
GANTENBEIN, Banjamin, Christian KRoPF, Carlo Rodolfo LARGIADER &
Adolf SCHOLL. Molecular and morphological evidence for the presence
of a new Buthid taxon (Scorpiones: Buthidae) on the Island of Cyprus
Pages
81-95
97-106
107-122
123-138
139-151
153-163
165-211
213232
REVUE SUISSE DE ZOOLOGIE
Volume 107 — Number 1
JUVARA-BALS, Ilinca & Vojciech WITALINSKI. Description of five new
species of Holoparasitus s. str. with redescription of H. apenninorum
(Berlese, 1906) and H. cultriger (Berlese, 1906) from Italy and Spain
(Sean Gamasıda-Barasıtidae).r or RR
SILVA, Carla Maria Menegola da & Beatriz MOTHES. Three new species of
Geodia Lamarck, 1815 (Porifera, Demospongiae) from the bathyal
depths off Brazilian coast, Southwestern Atlantic.................
MAHUNKA, Sandor & Luise MAHUNKA-Papp. Oribatids from Switzerland III
(Acari: Oribatida: Oppiidae | and Quadroppiidae). (Acarologica Ge-
Haven ar NEIL) wipes ewe oe nce: Ne alto cok ed er
GIL DE PERTIERRA, Alicia A. & Alain de CHAMBRIER. Rudolphiella szidati
sp. n. (Proteocephalidea: Monticelliidae, Rudolphiellinae) parasite of
Luciopimelodus pati (Valenciennes, 1840) (Pisces: Pimelodidae) from
Argentina with new observations on Rudolphiella lobosa (Riggenbach,
DS OS) EMEA O: IR CARTIER, et RUES RC NES, (REA eee
BENJAMIN, Suresh P. & Rudy JOCQUÉ. Two new species of the genus
Suffasia from Sri Lanka (Araneae: Zodariidae). . ................
LANG, Claude. Diversity of zoobenthos in 47 rivers of western Switzerland:
HEIISIENIITÄTLENIR RM IONI E
Vit, Stanislav. Contribution to the knowledge of Eucinetidae (Coleoptera). .
BANZIGER, Ruth. Spatio-temporal distribution of size classes and larval
instars of aquatic insects (Ephemeroptera, Trichoptera and Leopi-
doptera) in a Potamogeton pectinatus L. bed (Lake Geneva, Switzer-
lands ra LR US kien a er E AUTOMATE. De RARE
BESUCHET, Claude & Stanislav VIT. European species of the genus Nanop-
thalmus Motschulsky (Coleoptera, Scydmaenidae)...............
HEYD, Andreas & Wolfgang PFEIFFER. Sound production in catfish (Siluro-
idei, Ostariophysi, Teleostei) and its relationship to phylogeny and
fright means BOC MGA DIR ON SERIE EB SER
GANTENBEIN, Banjamin, Christian KROPF, Carlo Rodolfo LARGIADER &
Adolf SCHOLL. Molecular and morphological evidence for the presence
of anew Buthid taxon (Scorpiones: Buthidae) on the Island of Cyprus
Indexed in CURRENT CONTENTS, SCIENCE CITATION INDEX
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Volume 107 - Number 1 - 2000
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SWISS JOURNAL OF ZOOLOGY
REVUE SUISSE DE ZOOLOGIE
REVUE SUISSE DE ZOOLOGIE
TOME 107 — FASCICULE 2
Publication subventionnee par:
ACADEMIE SUISSE DES SCIENCES NATURELLES ASSN
VILLE DE GENEVE
SOCIETE SUISSE DE ZOOLOGIE
VOLKER MAHNERT
Directeur du Muséum d'histoire naturelle de Genéve
MANUEL RUEDI
Chargé de recherche au Muséum d'histoire naturelle de Genève
CHARLES LIENHARD
Charge de recherche au Muséum d’histoire naturelle de Geneve
Comité de lecture
Il est constitué en outre du président de la Société suisse de Zoologie, du directeur du
Muséum de Genève et de représentants des Instituts de zoologie des universités
suisses.
Les manuscrits sont soumis à des experts d'institutions suisses ou étrangères selon le
sujet étudié.
La préférence sera donnée aux travaux concernant les domaines suivants: biogéo-
graphie, systématique, évolution, écologie, éthologie, morphologie et anatomie
comparée, physiologie.
Administration
MUSÉUM D'HISTOIRE NATURELLE
1211 GENÈVE 6
Internet: http://www.ville-ge.ch/musinfo/mhng/page/rsz.htm
PRIX DE L'ABONNEMENT:
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(en francs suisses)
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| à la rédaction de la Revue suisse de Zoologie,
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REVUE SUISSE DE ZOOLOGIE 107 (2): 233-243; juin 2000
Etat trophique du lac de Morat indiqué par le zoobenthos:
tendance 1980-1998
Claude LANG
Conservation de la faune, Marquisat 1, CH-1025 St-Sulpice, Suisse.
E-mail: claude.lang @sffn.vd.ch
Trophic state of Lake Morat indicated by the zoobenthos: the 1980-
1998 trend. - Total phosphorus concentrations have strongly decreased
between 1982 and 1998 in the water of Lake Morat (Switzerland). In
response to this improvement, the numbers of tubificids (Tubifex tubifex
and Potamothrix hammoniensis mostly) decreased in the deepest area (40-
45 m), but the other species (Limnodrilus profundicola and L. hoffmeisteri)
present therein, remained scarce. In contrast at a depth of 20 m, chironomid
larvae (Chironomus and Procladius) and the tubificid Potamothrix molda-
viensis became more abundant in 1998. These changes indicated a slow
improvement of benthic conditions but, in 1998, the zoobenthos was
always characteristic for a eutrophic lake.
Key-words: chironomid — eutrophication — indicator — lake — oligochaete -
zoobenthos.
INTRODUCTION
Les concentrations en phosphore total dans l'eau du lac de Morat ont fortement
diminué en réponse aux mesures d’assainissement prises dans le bassin versant,
passant de 147 mg / m? en 1982 à 30 mg en 1998 (OPE, 1999). La baisse du phos-
phore devrait entrainer celle de la production des algues planctoniques, donc une
diminution de la sédimentation organique, et par conséquence une meilleure oxygé-
nation des couches d’eau profondes (Sas, 1989). La production des algues n'a pas été
mesurée mais les concentrations en oxygene ont augmente dans les eaux du fond
pendant la stagnation estivale (OPE, 1999). Cette amélioration de l'oxygénation
devrait faire sentir ses effets au niveau de la faune des sédiments profonds (faune
benthique ou zoobenthos). Celle-ci se compose principalement (si on se limite a la
macrofaune) d’oligochètes et de larves de chironomides (Diptera) dont les commu-
nautés d'espèces varient en fonction de l’état du lac et peuvent ainsi servir d’indi-
cateurs (Johnson et al., 1993).
Des conditions oligotrophes (peu de phosphore et beaucoup d'oxygène) pré-
valaient au début du siècle dans la plupart des grands lacs suisses (Sas, 1989). Les
nombres d’individus appartenant a des especes d’oligochetes et de chironomides
Manuscrit accepté le 26.01.2000
234 CLAUDE LANG
indicatrices de ce type de conditions ont diminué en réponse a l’augmentation du
phosphore entre 1960 et 1980, tant dans le Léman que dans le lac de Neuchatel (Lang,
1998, 1999). Lorsque ce nutriment a diminué, les nombres d’individus de ces espèces
ont augmenté à nouveau dans ces deux lacs, montrant ainsi que l’état des sédiments
profonds s’améliorait. Les espèces oligotrophes ne sont plus présentes dans la zone
profonde du lac de Morat où les espèces caractéristiques des lacs eutrophes consti-
tuent l’essentiel du zoobenthos entre 1980 et 1991 (Lang & Reymond, 1993).
En effet, ce lac est eutrophe depuis longtemps. En 1825 déjà (Jaag, 1948),
l’eau du lac de Morat était devenue rouge à cause de la prolifération d’une cyano-
bactérie (Oscillatoria rubescens, appelée maintenant Planktothrix rubescens DC).
Certains riverains, croyant que c'était le sang des soldats tués lors de la bataille de
Morat en 1476 qui remontait à la surface, ont appelé ce phénomène Sang des
Bourguignons. L’ apparition de cet organisme révèle que l’état d’un lac commence à
se dégrader (Sas, 1989). En 1936, la rareté de l’oxygène dans les couches d’eau
profondes pendant la stagnation estivale indique que les algues prolifèrent en surface;
le Sang des Bourguignons est toujours abondant, mais pas de façon continue (Rivier,
1936). La concentration en phosphore total qui est de 35 mg /m? en 1955, augmente
jusqu’en 1982 avant de commencer à redescendre (Liechti, 1989). En 1972, le lac est
qualifié d’eutrophe au vu de la composition chimique de ses sédiments (Davaud,
1976).
La présente étude cherche à évaluer et à interpréter la réponse du zoobenthos à
la baisse des concentrations en phosphore entre 1982 et 1998 en termes d’amélio-
ration de l’état biologique des sédiments profonds. En effet, le lac de Morat ne sera
restauré que lorsque ses sédiments le seront puisqu'ils reçoivent, sous une forme ou
une autre, tout ce qui est produit dans la colonne d’eau (Hakanson & Jansson, 1983).
Si l’état des sédiments s’améliore, la densité des oligochètes devrait diminuer tandis
que celle des larves de chironomides devrait augmenter (Wiederholm, 1980). De plus,
la composition des espèces devrait se modifier au sein de ces deux groupes: les
espèces très résistantes au manque d'oxygène devraient être remplacées peu à peu par
celles qui prévalaient avant la phase d’eutrophisation intense (Lang, 1998).
STATIONS ET MÉTHODES
Le lac de Morat qui couvre une surface de 23 km?, est relativement peu
profond (profondeurs moyenne et maximale: 23 m et 45 m respectivement) ce qui
explique sa sensibilité vis-à-vis de l’augmentation des concentrations en phosphore
(Liechti, 1989). Son principal affluent, la Broye, contribue pour 87 % à ses apports en
phosphore (OPE, 1999); c’est donc surtout l’état du bassin versant de cet affluent qui
détermine celui du lac. Les mesures d’assainissement prises ont entraîné la baisse
dans l’eau du lac des concentrations en phosphore total de 62 mg / m? en 1987 à 31
mg en 1996 (OPE, 1999). De ce fait, les concentrations moyennes en oxygène à 40 m
de profondeur entre les mois de juillet et de novembre ont passé de 1.8 mg /1 en 1987
à 3.6 mg / l en 1996; les valeurs extrêmes étant respectivement 1.5 à 2.9 mg et 2.1 à
4.9 mg. Les années antérieures à 1987 et postérieures à 1996 ne sont pas incluses dans
la comparaison parce que la fréquence des mesures d’oxygène n’y est suffisante ni
pour calculer une concentration moyenne pendant l’été ni pour estimer la durée de la
ZOOBENTHOS DU LAC DE MORAT 235
période où l’oxygene est rare. Pour cette raison, il n'est pas possible de comparer
directement l’évolution des concentrations en oxygène entre 1980 et 1998 a celle du
zoobenthos.
Entre 1980 et 1998, le zoobenthos du lac de Morat a été étudié dans 11 stations
de prélévements localisées entre 10 m et 45 m de profondeur (Fig. 1). A cela
s’ajoutent, en 1994, 42 stations réparties régulièrement sur l’ensemble du lac et, en
Môtier
Faoug I
o)
Fic. 1
Localisation des onze stations de prélèvements visitées entre 1980 et 1998 dans le lac de Morat.
Les flèches indiquent les points d’entrée et de sortie de la Broye, le principal affluent.
1998, 15 stations distantes de 200 m les unes des autres, placées sur un transect qui va
de la station 7 à la station 8. Dans chaque station, des carottes de sédiment de 16 cm?
chacune ont été prélevées en nombre variable selon les années et les profondeurs
(Tab. 1). Les prélèvements sont effectués au moyen d’un carottier, descendu depuis la
surface à l’extrémité d’un cable, dans la plupart des stations, sauf dans les stations 1 et
3, visitées en plongée par l’auteur en 1987 et en 1998. Seule la station 1 est située
dans la zone littorale (10 m), toutes les autres le sont dans la zone profonde (20 m à
45 m).
En laboratoire, le sédiment contenu dans chaque carotte est tamisé (ouverture
de maille: 0.2 mm). Le refus du tamis est fixé au formol 5 %. La faune benthique est
ensuite séparée du sédiment, sous une loupe lumineuse à l’aide de pinces fines. Les
236 CLAUDE LANG
TABLEAU |
Frequence et abondance (A) relatives (%) des especes de tubificides du lac de Morat en
fonction de la profondeur, de la station et de l’année. Fréquences calculées seulement à partir
des individus sexuellement matures. Fréquences 1980 pas comparables: + présence. Espece |
caracteristique des lacs mésotrophes; especes 2 a 7 caractéristiques des lacs eutrophes (Milbrink
1973):
Profondeur (m)
Especes 19: 10, 20.207220 20 203372402407 AO OISE
1. Potamothrix moldaviensis 50 83 IM
(Vejdovsky, Mrazek)
2. Ilyodrilus templetoni 33 20 6
(Southern)
3. Limnodrilus claparedeanus 17 6 7
Ratzel
4. Limnodrilus hoffmeisteri 33 GTS 6
Claparède
5. Limnodrilus profundicola 17 17 10 + £25 19 31 + 12
(Verrill)
6. Potamothrix hammoniensis 83 + 25 ..6 100° ASE 87 IS CO- -S
(Michaelsen)
7. Tubifex tubifex (Miller) 337 920i a = oe 81 44 53 70 76
Espece | (A) 1228019 2
Espèces 3 à 5 (A) ID a ZITTI 7 200001 2
Espèces 2, 6, 7 (A) 66,092. Sess) 7109 737,697 789293298 5999000
Année $7 98 78571807, 8491 98" OA 808 AI EI See Oo RS 6
Jour et mois 28.4 30-3) 877 B07 25:6925'6° 31.372167 30723.602 2031097238]
Nbre carottes (16 cm2) 67212772105 1677 116 7167, 22 SRO IS 30 2021
Nbre vers identifiés 40 48 24 121 125 59 56 402 169 306 143 134 170 174
Stations (fig. 1) ] | 10) 2-5. 2-5), 9225) 3) Fig23)}6=9) 16-9) 6-9 Sl
oligochetes, les larves de chironomides et de Chaoborus sont comptés et pesés
(biomasse), apres passage sur du papier buvard pour enlever l’eau en exces. Tous les
tubificidés (ou un sous-échantillon s'ils sont trop nombreux), dont le diamètre a l’état
fixe dépasse 0.29 mm (Lang, 1998), sont montes entre lame et lamelle dans un milieu
éclaircissant (Reymond, 1994) afin d’être identifié sous le microscope. Seuls les
individus sexuellement matures des taxons présents dans le lac de Morat peuvent être
identifies jusqu’au niveau de l’espece. Cependant, les immatures peuvent tout de
méme étre classés dans des groupes d’especes dont la valeur indicatrice est analogue
(Lang, 1998). Nous distinguons ainsi les especes caractéristiques des lacs oligo-
trophes, mésotrophes ou eutrophes. La fréquence relative d’une espece s’exprime
comme le pourcentage de carottes dans lesquelles cette espèce est présente. L’abon-
dance relative d’une espèce se définit comme le nombre d’individus de cette espèce
rapporté, sous forme de pourcentage, au nombre total d'individus identifiés dans une
carotte ou un ensemble de carottes.
De 1984 à 1998, les résultats sont analysés carotte par carotte, contrairement à
1980, où les 4 carottes prises dans chacune des 8 stations visitées cette année-là sont
cumulées. De plus, la biomasse n’a pas été mesurée en 1980. Pour ces raisons, les
ZOOBENTHOS DU LAC DE MORAT 237
résultats de 1980 ne peuvent pas étre inclus dans toutes les comparaisons effectuées
entre les années. Les seules campagnes de prélevements vraiment comparables entre
elles à tous les points de vue sont celles effectuées en 1984, 1991 et en 1998 a 20 m et
40 m de profondeur.
La biomasse du zoobenthos (g/m?) peut être calculée à partir des concen-
trations en phosphore total (mg/m?) en utilisant une relation empirique établie dans
d’autres lacs (Hanson & Peters, 1984):
Log; biomasse = 0.708 log} phosphore + 0.092
La concentration moyenne du phosphore, calculée sur les cinq années précédant le
prélèvement du zoobenthos, est utilisée dans cette expression pour tenir compte du
fait que le zoobenthos réagit souvent avec un temps de latence à la baisse du
phosphore (Lang, 1998). Si la biomasse observée est proche de la biomasse calculée,
cela signifie qu'il n’y a pas de décalage entre l’état trophique du sédiment et celui de
l’eau. Ce type de relation empirique permet de replacer l’évolution trophique d’un lac
particulier dans un contexte plus général et de mettre ainsi en évidence d'éventuelles
anomalies (Lang, 1998).
RÉSULTATS
Dans le lac de Morat, la diversité du zoobenthos diminue avec la profondeur
(Tab. 1) ce qui reflète la baisse des concentrations en oxygène (OPE, 1999). A 40 m
et 45 m de fond, le zoobenthos se compose presque exclusivement de deux espèces de
tubificidés, très résistantes au manque d'oxygène, dont la dominance numérique
caractérise les lacs eutrophes: Potamothrix hammoniensis et Tubifex tubifex (Mil-
brink, 1973). Le pourcentage de ces deux espèces dans les communautés d’oligo-
chètes ne change guère à 40 m entre 1980 et 1998. A 20 m de profondeur, le pourcen-
tage des espèces du genre Limnodrilus augmente par rapport à 40 m, mais reste assez
constant entre 1980 et 1998. Notons en 1998 la présence à 20 m de Potamothrix
moldaviensis, une espèce plutôt caractéristique des lacs mésotrophes (Milbrink,
1973), qui est relativement abondante à 10 m, tant en 1987 qu’en 1998. Les cinq
premières espèces du tableau 1 sont présentes dans le Léman jusqu’à 150 m de pro-
fondeur au moins (Lang, 1998), ce qui montre que ce n’est pas ce facteur qui limite
leur distribution verticale dans le lac de Morat, mais plutôt le manque d'oxygène.
A 40 m de profondeur, la biomasse du zoobenthos ne change pas entre 1984 et
1998 (Tab. 2), mais le nombre de tubificidés diminue significativement (test de
Kruskal Wallis: P = 0.084 et P = 0.007 respectivement). À 20 m de fond, la biomasse
du zoobenthos augmente entre 1984 et 1998. Cette augmentation est surtout le fait des
larves de chironomides qui constituent en 1998 le 54 % de la biomasse. Seuls les
genres Chironomus et Procladius, résistants au manque d’oxygène (Brinkhurst,
1974), sont présents et leur abondance est la même (1500 individus/m?). Une ten-
dance semblable s’observe en1998 à 10 m de profondeur: les chironomides forment le
43 % de la biomasse, Chironomus et Procladius sont dominants. Mis à part 1998 (à
10 m et 20 m), les chironomides ne représentent qu’une petite partie de la biomasse
totale dans la zone profonde du lac de Morat qui se caractérise par la prédominance
des tubificidés. Notons enfin la présence de quelques larves de Chaoborus à 40 m de
profondeur.
238 CLAUDE LANG
TABLEAU 2
Biomasse moyenne ( g / m2) du zoobenthos et nombre moyen d’oligochètes par m? en fonction
de la profondeur et de l’année dans cing lacs: Morat, Léman, Neuchatel, Joux, Bret. Sources:
cette étude, Lang 1998 et 1999, résultats non publiés.
Biomasse 2) Nombre
Profondeur ST
Lac (m) Année nl) Moy. ESM Max. Moy. ESM Max.
Morat 10 1987 6 13.5 3.1 24 30104 3016 39375
1998 12 24.1 3.2 38 12604 1873 27500
20 1980 16 - 3) = = 23750 à =
1984 16 9.3 2.6 45 18085 2374 41250
1991 16 7.8 1.5 19 11289 1341 25000
1998 12 49.7 5.4 82 30677 3439 56875
35 1994 42 43.2 8.5 356 49405 6074 245625
40 1980 16 a = = 29375 = È
1984 16 22.8 2.9 56 26875 3159 62500
199] 16 19.9 3.9 66 21679 3408 55000
1998 30 211 4.8 108 21541 5079 125625
45 1985 20 33.1 6.5 119 21312 3212 53750
1986 21 19.3 3.2 62 20833 2587 52500
Léman 40 199] 192 24.0 1.6 219 24687 2111 274375
1996 170 17.3 0.6 47 12092 585, 248105
60 1994 159 13.5 0.6 38 20243 929 58750
1999 159 11.6 0.7 57 12448 WS NHS
Neuchatel 40 1992 171 10.2 0.5 35 9181 570 61250
1997 175 9.4 0.5 32 8546 569 45625
Joux 25 1998 450 52.1 1.5 186 39164 1001 230000
Bret 20 1998 35 21:8 2.3 52 929 202 4375
Leman 270 1998 25 4.0 0.8 19 5500 686 12500
1) nbre de carottes de 16 cm?
2) Moy. moyenne, ESM erreur standard de la moyenne, Max. valeur maximale.
3) Résultats manquent.
La campagne de prélévements effectuée en 1994 montre (Fig. 2 et 3) la
répartition et la composition du zoobenthos dans l’ensemble la zone profonde du lac
de Morat. Les valeurs de la biomasse du zoobenthos qui sont plus élevées dans la
zone du lac influencée directement par les apports organiques de la Broye, le principal
affluent (70 % des apports en eau, Liechti, 1989), diminuent en général au fur et a
mesure que l’on s’en éloigne, tout comme les taux de sédimentation organique
(Davaud, 1976). Tubifex tubifex et Potamothrix hammoniensis constituent l’essentiel
ZOOBENTHOS DU LAC DE MORAT 239
tant des tubificides (89 % des individus en moyenne) que du zoobenthos. Les autres
especes de tubificidés du genre Limnodrilus se rencontrent surtout a la périphérie de
la zone étudiée où la profondeur diminue. Les larves de chironomides ne sont
présentes que dans 6 des 42 stations visitées et, en nombre d’individus (un individu
seulement dans chacune des 6 stations), elles ne représentent que le 0.2 % du nombre
des tubificidés. Cependant, comme les prélèvements ont été effectués le 14 et le 21
juin, donc à la fin de la période d’émergence des chironomides, l’abondance des
larves est probablement sous-estimée. Des prélèvements effectués en avril seraient
plus représentatifs à cet égard.
569 570 571 572 573 574 575 576
FIG. 2
Répartition géographique de la biomasse du zoobenthos (g /m?) dans le lac de Morat en 1994.
Les coordonnées géographiques de la carte nationale de la Suisse sont indiquées en x et en y.
La biomasse moyenne du zoobenthos est de 43.2 g / m? dans les 42 stations
visitées en 1994 (Fig. 2); la biomasse médiane qui ne prend pas en compte les stations
très influencées par les apports de la Broye, est de 32.2 g / m?. Ces deux valeurs
observées sont bien supérieures à 16.5 g / m?, la biomasse calculée à partir de la
moyenne (39 mg / m?) des concentrations en phosphore dans l’eau entre 1989 et 1993
(voir Stations et méthodes). Par contre, la biomasse médiane observée en 1994 est très
proche de 35.2 g / m?, la biomasse calculée à partir de la moyenne des concentrations
en phosphore mesurées entre 1982 et 1986 (113.4 mg / m?). Cette différence montre
que la biomasse du zoobenthos réagit lentement à la baisse du phosphore si bien que
240 CLAUDE LANG
199.5
10074722100210033285
199.0 - D a Oo o D DO
33. 66 710071007100310053
198.5 - o o o o o o o
73 64 100 100 100 100 100 100
198.0 - [n] [m] je} D je} O D D
| 100 100 100 100 100 67 100 100
197.5 D D D Oo o D D D
62 94 100 100 8 71100
197.0- (a) D D Oo [=] [n] [a]
Broye
100 100 91 70 100 100
196.5 + o o o o o o
196.0 | U T T T T
569 570 571 572 57S 574 575 576
FIG. 3
Répartition géographique du pourcentage de Tubifex tubifex et de Potamothrix hammoniensis
dans les communautes de tubificidés du lac de Morat en 1994.
les sédiments restent capables en 1994 de nourrir des populations benthiques qui
correspondent encore au niveau de productivité observé dix ans plus tot dans l’eau du
lac.
DISCUSSION
Le lac de Morat est un des premiers lacs suisses dont l’état s’est dégradé sous
l’influence des activités humaines (Jaag, 1948). La prolifération du Sang des
Bourguignons en 1825 indique le début de cette dégradation. Une intensification des
activités agricoles dans le bassin versant de la Broye, le principal affluent du lac,
pourrait en avoir été la cause, comme c’est le cas depuis le moyen age déja pour de
nombreux lacs d’Europe (Anderson, 1995). Signalons par exemple la culture du tabac
qui s’est développée entre 1719 et 1819 dans la plaine de la Broye (Chuard &
Dessemontet, 1972 ). De même, la production de blé a augmenté de 38 % entre 1707-
1709 et 1787-1789 dans les balliages de Payerne et d’Avenches qui englobaient la
partie inferieure du bassin versant (Chevallaz, 1949). Cet auteur fait également
remarquer que l’agriculture vaudoise s’est fortement développée a partir de 1803
grace à la suppression des usages et des structures contraignants de l’ Ancien Régime.
ZOOBENTHOS DU LAC DE MORAT 241
Cette evolution rapide des pratiques agricoles s’effectue dans un contexte
météorologique particulier. Les années 1816 et 1817 se caractérisent en effet par des
pluies et des inondations exceptionnelles (Pfister, 1985). Celles-ci résultent du
bouleversement climatique engendré par la présence, dans la haute atmosphere, des
poussières provenant de l’éruption en 1815 du volcan indonésien Tambora (Whyte,
1995). Ces pluies, tombant sur les sols du bassin versant du lac de Morat, cultivés de
façon plus intensive, en ont probablement lessivé les nutriments, créant ainsi les
conditions qui ont provoqué en 1825 la prolifération du Sang des Bourguignons. Cette
longue histoire de dégradation (plus de 150 ans) laisse supposer que la restauration
biologique du lac de Morat prendra plus de temps que celle du Léman ou du lac de
Neuchatel ou la phase de détérioration n’a duré qu’une trentaine d’années (Lang,
1998, 1999).
Malgré la baisse spectaculaire des concentrations en phosphore dans l’eau du
lac de Morat, la composition du phytoplancton et son abondance, ainsi que celles du
périphyton épilithique, restent caractéristiques d’un lac eutrophe en 1988 et en 1989
(Reymond & Straub, 1993). De même, le zoobenthos ne se modifie pas de façon
fondamentale entre 1980 et 1998, tout au moins dans la zone profonde, seule étudiée
en détails. Les deux espèces de tubificidés qui restent les plus abondantes entre 1980
et 1998, Tubifex tubifex et Potamothrix hammoniensis, étaient déjà les seules
présentes en 1935 dans la zone la plus profonde du lac (Rivier, 1936). A cette époque,
elles colonisaient en petit nombre le fond du lac en compagnie des larves de
Chaoborus qui étaient relativement abondantes. Ce genre de faune se rencontre dans
des lacs très eutrophes, où le manque d’oxygene persiste pendant des mois. Dans ce
type de situation, l'abondance des tubificidés qui vivent constamment en contact avec
le fond du lac, diminue davantage que celle des Chaoborus qui peuvent migrer
chaque nuit vers la surface et les couches oxygénées (Brinkhurst, 1974). C’est la
situation observée en 1998 dans le lac de Bret à 20 m de profondeur où Chaoborus
constitue le 98 % de la biomasse (Tab. 2). Lorsque le lac est profond, les Chaoborus
sont absents: dans le Léman par exemple, seuls les tubificidés sont présents entre 270
m et 300 m de fond et leur biomasse est faible à cause du manque d’oxgene qui
caractérise cette zone (Lang, 1998). En 1972 comme en 1936, les sédiments profonds
du lac de Morat semblaient presque dépourvus de faune, mis à part les Chaoborus et
quelques tubificidés (Davaud, 1976). Entre 1980 et 1998, les tubificidés sont devenus
abondants et les Chaoborus rares, ce qui tendrait à montrer que l’état biologique des
sédiments s’est amélioré par rapport à la situation précédente. Toutefois, comme
celle-ci n’est pas décrite de façon quantitative dans les travaux de Rivier et de
Davaud, l'interprétation de la tendance peut être mise en doute.
Lorsque l’état d’un lac s'améliore après un épisode d’eutrophisation, la densité
des larves de chironomides augmente tandis que celle des oligochètes diminue
(Wiederholm, 1980). Cette tendance s’observe en 1998 dans le lac de Morat: à 10 m
et à 20 m de profondeur en ce qui concerne les chironomides, à 40 m en ce qui
concerne les tubificidés. Remarquons toutefois que l’abondance moyenne des oligo-
chètes en 1998 à 40 m est plus du double de celles observées à la même profondeur
dans le lac de Neuchâtel en 1992 et 1997 (Tab. 2); les valeurs du Léman à 40 m et à
60 m de profondeur se situent entre ces deux extrêmes. Cette séquence des abon-
dances — Morat, Léman, Neuchâtel — reflète la décroissance des concentrations en
242 CLAUDE LANG
phosphore entre ces trois lacs. Cependant, il existe des exceptions à cette règle. Dans
le lac de Joux par exemple, malgré la baisse du phosphore, la biomasse du zoobenthos
et l’abondance des tubificidés restent élevées à cause de la présence presque continue
du Sang des Bourguignons qui sédimente en masse sur le fond du lac entre 1980 et
1998. Une fois installé dans un lac, cet organisme peut en effet s’y maintenir malgré
les mesures d’assainissement prises (Sas, 1989). Son retour en force dans le lac de
Morat n’est d'ailleurs pas exclu.
L’amélioration constatée dans le lac de Morat est plus marquée à faible qu’à
grande profondeur. De ce fait, des prélèvements de zoobenthos concentrés sur les
bords de la zone profonde (entre 15 m et 30 m) auraient probablement mieux mis en
évidence la restauration de l’état des sédiments que le programme suivi. Une autre
strategie possible consiste a augmenter l’intervalle de temps entre les campagnes de
prélevements de maniere a ce que le zoobenthos puisse répondre de facon plus claire
à la baisse du phosphore. Par exemple, la campagne de 1994 qui couvre l’ensemble de
la zone profonde (Fig. 2 et 3), pourrait étre refaite en 2004, si possible en avril avant
l’émergence des chironomides. Si ces dix ans d’amélioration chimique ont eu un effet
positif au niveau du sédiment, les larves de chironomides devraient devenir plus
nombreuses et recoloniser l’ensemble de la zone profonde, la baisse de l’abondance
des tubificidés devrait se poursuivre et la dominance de Tubifex tubifex et Potamo-
thrix hammoniensis devrait diminuer au profit d’autres espèces moins résistantes au
manque d'oxygène. La colonisation par Potamothrix moldaviensis, espèce indicatrice
de conditions mésotrophes, des sédiments situés à 45 m de profondeur montrerait que
le lac de Morat est vraiment en voie de restauration.
Si la baisse du phosphore se poursuit, le retour des espèces indicatrices de
conditions oligotrophes, comme dans le Léman et le lac de Neuchâtel (Lang, 1998,
1999), n’est pas exclu à long terme. L’exemple du lac Esrom montre que celles-ci
peuvent en effet rester présentes à très faible profondeur (entre 0.5 m et 1.5 m) dans le
littoral d’un lac eutrophe, comme c’est le cas du lumbriculidé Srylodrilus heringianus
(Dall et al., 1990). Signalons à ce propos qu’en 1905 Spirosperma ferox et Psammo-
ryctides barbatus, deux espèces de tubificidés indicatrices de conditions oligo-
mésotrophes (Milbrink, 1973), étaient présentes à 4 m de profondeur dans le lac de
Morat (Piguet, 1906). Si elles ont subsisté à très faible profondeur comme c’est
probable, elles pourraient graduellement recoloniser le lac au fur et à mesure des
progrès de la restauration.
REMERCIEMENTS
L’aide d'Olivier Reymond en laboratoire et celle de Raymond Ducret sur le
terrain m'ont permis de mener à bien ce travail. Les remarques du comité de lecture
ont contribué à améliorer cet article.
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REVUE SUISSE DE ZOOLOGIE 107 (2): 245-257; juin 2000
A new species of Astyanax (Characiformes, Characidae)
from Uruguay river basin in Argentina, with remarks
on hook presence in Characidae
Maria de las Mercedes AZPELICUETA! and José O. GARCIA?
| Dep. Cientifico Zoologia de Vertebrados, Fac. de Ciencias Naturales y Museo de
La Plata, Paseo del Bosque, 1900 La Plata, Argentina.
? Fac. de Cs. Exactas y Naturales, Universidad Nacional de Misiones,
Rivadavia 2370, 3300 Posadas, Argentina.
A new species of Astyanax (Characiformes, Characidae) from Uruguay
river basin in Argentina, with remarks on hook presence in Chara-
cidae. - In the present paper the new species Astyanax ojiara sp. n. is
described, from the headwaters of Yaboti river, an affluent of Uruguay river
in the province of Misiones, Argentinean northeast. A combination of
characters differentiates the new species from other congeners: one
heptacuspid maxillary teeth; teeth of inner premaxillary row gently expan-
ded distally; 7-9 dentary teeth decreasing in size anteroposteriorly; males
with hooks in all fins; large males with pelvic axillary scale bearing | to 8
hooks; 36-38 perforated scales in lateral line; 20-23 branched anal-fin rays.
Number of scales in the lateral line and branched anal-fin rays, humeral
spot, and low body of Astyanax ojiara sp. n. resemble those of A. eigen-
manniorum, but maxillary, premaxillary and dentary teeth differ in both
species. The disposition of the dentary teeth is also similar in A. taeniatus
and A. giton but A. ojiara sp. n. has one heptacuspid maxillary tooth.
Key-words: Characiformes - Characidae - Astvanax - new species -
Uruguay river.
INTRODUCTION
In the last years, the genus Astyanax was not revised and an accurate definition
of the genus is still pendant. Eigenmann (1921, 1927) analised the genus and, sub-
sequently, many authors followed his results. Géry (1977) provided an arrangement of
the species in groups, and identification keys. Recently, Zanata (1997) erected a new
genus - Jupiaba - for many species previously known as Astyanax or Deuterodon.
Jupiaba is defined by the presence of a diagnostic pelvic bone, with unique disposition
of muscles. The taxonomic status of some species of the genus Astyanax is not comple-
tely clear and the interrelationships of the species remain unresolved. A detailed
revision of the remaining nominal species of Astyanax is necessary.
Manuscript accepted 27.12.1999
246 MARIA DE LAS MERCEDES AZPELICUETA & JOSE O. GARCIA
In the rio de la Plata basin, about twenty nominal species of Astyanax have been
recorded, although the records of some species are doubtful. In the southwest of the
Brazilian shield, the main course of the rio Uruguay flows from East to West, turning to
the southwest and ending in the rio de la Plata. Some affluents of the river originate
from the Sierra de Misiones, which reach about 800 m a.s.l., and presently constitute an
effective barrier between waters of the Uruguay and Paranä rivers, in northeastern
Argentina. The arroyo Yaboti comes from the highest area of Sierra de Misiones; in one
of its headwaters, the arroyo Benitez, a new species of Astyanax was collected in the
vicinity of the city of San Pedro. The description of the new species is presented in this
paper.
MATERIAL AND METHODS
The specimens examined in this study were cleared and counterstained (C&S)
following Taylor & Van Dyke (1985). Measurements are straight distances taken with
calliper to nearest 0.1 mm. Material is deposited in the Field Museum of Natural
History, Chicago (FMNH); Muséum d’histoire naturelle de Geneve, Geneve (MHNG);
Facultad de Ciencias Naturales y Museo de La Plata, La Plata (MLP); Museu de
Zoologia da Universidade de Sao Paulo, Sao Paulo (MZUSP).
Comparative material (SL in mm). Astyanax eigenmanniorum (Cope, 1876):
MLP 5202, 5 ex., 56.5-68.5, Argentina, Cordoba, rio Primero frente a Capilla de los
Remedios; MLP 9160, 6 ex., 36.8-80.2, Argentina, Buenos Aires, Los Talas. Cleared
and stained material: Personal collection of MA. Astyanax abramis (Jenyns, 1842): 2
ex., 74.5-92.0, Argentina, Buenos Aires, rio de la Plata en Punta Lara; 2 ex., 80.6-98.8,
Argentina, Misiones, rio Piray-Mini. Astyanax alburnus (Hensel, 1870): 5 ex., 40.5-
47.2 , Uruguay, rio Yaguarön. Astyanax alleni (Eigenmann and McAtee, 1907): 2 ex.,
62.4-72.9, Argentina, Corrientes, rio Riachuelo; 1 ex., 66.0, same collecting data.
Astyanax cf. asuncionensis Géry, 1972: 2 ex., 28.0-37.6, Argentina, Santa Fe, Isla Los
Sapos; 2 ex., 80.4-92.7, Argentina, Misiones, rio Uruguay en San Isidro. Astyanax
eigenmanniorum: 2 ex., 17.7-33.0, Argentina, Buenos Aires, desembocadura del rio
Colorado; 2 ex., 28.0-30.5, Argentina, Buenos Aires, Laguna de Gomez; 1 ex., 60.3,
Argentina, Buenos Aires, rio de la Plata; 4 ex., 51.5-82.1, Argentina, Misiones, arroyo
Piray-Mini; 1 ex., 45.0, Brasil, Rio Grande do Sul, Viamäo, agude Charolés; Astyanax
cf. fasciatus (Cuvier, 1819): 2 ex., 91.0-106.5, Argentina, Misiones, rio Uruguay en San
Isidro. Astyanax ojiara sp. n.: 3 females, 46.2-63.0; 7 males, 37.8-58.0, Argentina,
Misiones, arroyo Benitez.
TAXONOMY
Astyanax ojiara Sp. n. Figs 1-13, table 1
Holotype (Fig. 1). MLP 9470, male, 50.5 mm SL, Argentina, province of Misiones,
arroyo Benitez, headwaters of rio Yaboty, an affluent of rio Uruguay, coll. J. O. Garcia, May
1983.
Paratypes. (Collecting data as holotype). MLP 9471, 2 males, 38.0-45.0 mm. MLP 9472,
12 females, 39.8-72.0 mm. MHNG 2605.67, 10 males (measured); MHNG 2606.35, 3 females,
44.0-53.5 mm, (not measured). FMNH 98319, 5 ex. MZUSP 40255, 5 ex.
A NEW SPECIES OF ASTYANAX FROM ARGENTINA 247
DIAGNOSIS
The species is distinguished by a combination of characters: one maxillary teeth
with seven small cusps; teeth of inner premaxillary row gently expanded distally; 7-9
dentary teeth decreasing in size anteroposteriorly; large males with hooks ın all fins;
large males with | to 8 hooks in the pelvic axillary scale; 36-38 perforated scales in the
lateral series; and iv-v,20-23 anal-fin rays. Also, presence of humeral spot vertically
elongated, a second faint humeral spot, caudal spot continued on middle caudal rays,
and low body depth help to differentiate the new species.
DESCRIPTION
Morphometrics of holotype and 24 paratypes are presented in table 1. Astyanax
with low body (Fig. 1), maximum body depth at dorsal-fin origin. Dorsal profile of
body slightly convex from snout to posterior tip of supraoccipital process, angled
behind supraoccipital, strongly marked in medium sized and large females. Dorsal
profile of body gently curved from this point to origin of dorsal fin; in large females,
that portion almost straight; slanted ventrally from dorsal-fin origin to caudal peduncle.
Dorsal profile of caudal peduncle straight; ventral profile slightly convex or straight.
Ventral profile of body slightly curved from tip of snout to pelvic-fin origin, straight
between this point and anal-fin origin, and slanted dorsally to caudal peduncle. Body
rounded between pectoral and pelvic fins. Body laterally compressed between pelvic
and anal fins.
Dorsal-fin origin nearer tip of snout than base of caudal-fin rays, equally distant
in some small specimens. Pelvic-fin origin anterior to vertical through dorsal fin-origin.
Adipose fin anterior to bases of last branched anal-fin rays. Tip of pectoral fin falling
near, sometimes surpassing, pelvic-fin origin; in large females, tip of pectoral fin far
from that origin. Tip of pelvic fin reaching anal-fin origin in males; in females, pelvic-
fin tip far from that origin, even in many small specimens.
Dorsal fin iii, 9; posterior margin of dorsal fin slightly rounded, first branched
dorsal-fin ray longest. In males, all dorsal-fin rays, excluded last one and unbranched
rays, with slender hooks, directed outward, most of them curved ventrally, more
abundant on posterior branch of each ray (Fig. 2), usually, one pair on each segment.
Few hooks developed on ray tips of small specimens.
Anal-fin iv-v, 20-23 (holotype 21). Posterior margin almost straight in males; in
females, first five branched rays produced forming a small lobe. Anal fin of all males
bearing hooks directed posteriorly and outward, slightly curved dorsally. Large
specimens with hooks on last branched anal-fin ray and also on largest unbranched ray.
Hooks placed on all branches of the ray (Fig. 3), sometimes two or three pairs on each
segment.
Caudal fin bearing 10-12 dorsal and 9-10 ventral procurrent rays; one
unbranched and 9 branched principal rays in upper lobe; 8 branched and 1 unbranched
principal rays in lower lobe. Usually, caudal lobes similar in size; sometimes, lower one
slightly longer. Few hooks occurring on distal tips of middle caudal-fin rays in males
(Fig. 4). In some specimens, few hooks scattered on all fin rays, included principal
unbranched one. Hooks directed outward, most of them on lower lobe curved dorsally
248 MARIA DE LAS MERCEDES AZPELICUETA & JOSÉ O. GARCIA
FIG. 1
Astyanax ojiara new species, holotype, MLP 9470, male, SL 50.5 mm, Argentina, province of
Misiones, arroyo Benitez, headwaters of rio Yaboty, an affluent of rio Uruguay.
and those ones of upper lobe curved ventrally. Some small specimens with 2 or 3 pairs
of hooks only; one 43.0 mm SL male without hooks.
Pectoral-fin 1, 10-13 (holotype i, 12), rays bearing hooks of different size in
males. In medium sized and large specimens hooks developed on first unbranched ray
and until 9 branched rays. Sometimes more than one pair of hooks on each segment,
some of them directed dorsally and other ones ventrally, most of them directed inward
(Fig. 5). Very few hooks developed in small specimens.
Pelvic fin i, 7; hooks developed on branched rays, occasionally on first
unbranched ray also in large specimens (Fig. 6). One or two pairs of hooks on each
segment, in different branches of ray, most of them directed inward, slightly curved
anteriorly. Usually, one ray with scarce hooks in small specimens.
Head length moderate, mouth terminal, horizontal; snout short. Lower jaw
slightly longer. Premaxilla with short ascending process, bearing two series of teeth.
Usually, 3 pentacuspid or tricuspid teeth in outer series, sometimes two or four teeth,
central cusp larger. Inner series of premaxillary teeth consisting of 5 teeth, gently
expanded distally, slightly compressed at distal tips (Fig. 8). Symphysial tooth narrower
and deeper, with 4 or 6 cusps. Second tooth widest, with 7 cusps. Third and fourth teeth
with 6-7 cusps. Fifth tooth smaller, with 5 cusps. In all teeth, central cusp slightly larger
than remaining ones. Maxilla short, scarcely lobed posteriorly, almost reaching or
scarcely surpassing vertical through anterior orbital margin. One compressed maxillary
tooth, with seven small cusps (Fig. 9). Few specimens (about 8 %) with two maxillary
teeth, one of them heptacuspid. Dentary bearing 7 to 9 (usually 8) teeth with broad
bases, decreasing in size anteroposteriorly. Symphysial tooth narrower, with 5-6 cusps.
Second tooth widest. Second, third and fourth teeth with 5-7 cusps. Fifth tooth with 3-5
A NEW SPECIES OF ASTYANAX FROM ARGENTINA 249
Fics 2-4
Astyanax ojiara, left view. 2: dorsal fin, SL 44.5 mm, scale= 1 mm; 3: detail of hooks on third
branched anal-fin ray, SL 39.6 mm, scale= 0.5 mm; 4: detail of hooks on middle caudal-fin rays,
SL 39.6 mm, scale= 1 mm.
250 MARIA DE LAS MERCEDES AZPELICUETA & JOSE O. GARCIA
cusps, sixth and seventh teeth with 1 to 3 cusps; eighth tooth - and nineth one when
present - always conic (Fig. 10).
Eye small, interorbital wide. Six infraorbitals well developed; third infraorbital
almost contacting sensory tube of preopercle. Anterior fontanelle triangular, widening
posteriorly; posterior fontanel long, extending onto supraoccipital process base.
Scales cycloid, crenate. Lateral series with 36-38 perforated scales (holotype
with 37, one specimen with 39). Lateral line running on lower half of caudal peduncle,
ending in a long tube without lamina, between caudal rays. Five scales between dorsal-
fin origin and lateral line; 5 between lateral line and ventral-fin origin. Thirteen or
fourteen scales around caudal peduncle. Eleven or twelve scales forming a regular row
between supraoccipital process and dorsal-fin origin; sometimes, 14 scales in an
irregular row. Ten to twelve (until 14) rectangular scales placed on anal-fin base,
covering all unbranched and twelve branched anal-fin rays. Scales placed on basal fifth
of caudal lobes. A narrow axillary scale present dorsal to pelvic-fin insertion, oval, as
long as one third of pelvic fin sometimes. Axillary scale bearing two hooks in its
posterior inner area in medium and large males; although, a high number of hooks
developed (8 in largest male, 56 mm SL, Figs 11-13). Young males with a ridge in that
area of scale.
In ten cleared and stained specimens, first arch bearing 17-19 gill-rakers: 2 on
hypobranchial, 8 on ceratobranchial, 1 on cartilage, and 6 to 8 on epibranchial.
Vertebral counts including Weberian apparatus and CUI+PUI as one element: 32 (1
sp.), 33 (2 sp.), and 34 (7 sp.).
Coloration of alcohol preserved specimens: Background light brown, dorsal
region of flanks and head darker. A humeral spot well developed, dorsoventrally
expanded, at level of pectoral-fin origin. A second lateral spot faint and smaller, usually
rounded. A dark lateral band with different intensity in coloration and width crossing
flanks. A deep and very narrow line of chromatophores runnig from humeral spot to
base of middle caudal-fin rays. Most specimens with dark chromatophores on inner
opercular surface forming a spot.
Dark chromatophores on distal margin of anal fin, forming a band; two or three
first dorsal rays with black chromatophores; middle caudal-fin rays black, also tips of
caudal-fin rays with dark chromatophores; pectoral and pelvic fins hyaline.
ETYMOLOGY
The specific epithet is the name of a spirit, protector of the fresh waters, in the
tupi-guarani language.
DISCUSSION
Eigenmann (1921, 1927) separated the species of the genus Astyanax into three
subgenera that were maintained by many authors. Astyanax ojiara has low body depth
and a complete series of scales in the predorsal area. Following Géry (1977), A. ojiara
is placed in the group with 32-41 scales in the lateral line. Among those species, the
number of anal-fin rays of A. ojiara agrees with that of A. fasciatus group. Like Géry
(1977) pointed out, the other group is defined by the presence of 17-24 anal-fin rays,
A NEW SPECIES OF ASTYANAX FROM ARGENTINA 25]
GE
=
er 9
Za
N
EI
es Ss Ee
7
a
Fics 5-7
Astyanax ojiara, dorsal view, SL 45.0 mm. 5: left pectoral fin, scale= 1 mm; 6: detail of hooks on
second branched pelvic-fin ray, scale= 0.5 mm. 7: A. eigenmanniorum, dorsal view, SL 45.0 mm,
detail of the same ray, scale= 0.5 mm.
252 MARIA DE LAS MERCEDES AZPELICUETA & JOSE O. GARCIA
except A. eigenmanniorum. The number of perforated scales in the lateral line and the
number of anal-fin rays of A. ojiara ressemble those of A. eigenmanniorum.
Nonetheless, the number and shape of teeth greatly differ. Astyanax eigenmanniorum
has 4-5 teeth in the outer series of premaxilla and 5 teeth in the inner one, with the
central cusp notably larger; the maxillary tooth has 3 to 5 cusps (Azpelicueta, 1979); all
those teeth always have broad bases. The dentary bears 4 large teeth, a median one, and
a series with 5 to 7 very small teeth. In comparison, the maxilla of A. eigenmanniorum
is narrow and long, always reaching the anterior third of the eye; the premaxilla has a
longer dorsal process. Astyanax eigenmanniorum, as described by Cope (1876), has
pectoral fins that reach ventral-fin origin, a character always present in the Brazilian
specimens examined; the length of pelvic fins and the preanal distance have similar
values that those of males of A. ojiara. The pelvic-fin hooks of A. eigenmanniorum are
strong and regularly placed (Figs 6, 7); the anal-fin hooks develop on the first ten rays
whereas they are smaller, more numerous and are present on many anal-fin rays of A.
ojiara. In A. eigenmanniorum, the eye, the postorbital length, and the interorbital
distance are larger.
From other species of the A. eigenmanniorum group, A. ojiara is distinguished
by one maxillary heptacuspid tooth, and the hooks developed in all fins of males.
Astyanax taeniatus (Jenyns, 1842) and A. giton Eigenmann, 1908 have dentary teeth
decreasing in size anteroposteriorly. Lucena & Lucena (1992) discussed the presence of
that feature as a character that evolved in different species of the genera Deuterodon
and Astyanax.
Different measurement relations in both sexes of A. ojiara are the length of
pelvic fin, the distances between last branched anal-fin ray-hypural joint, snout tip-anal-
fin origin, and origins of pelvic and anal fins (Table 1). Two thirds of the sample
examined were males.
Presence of hooks in Characidae
Sexual dimorphism appears in many characids; hooks develop in different fins
of males, excluding the genus Cheirodon in which female pelvic fins bear hooks
(Malabarba, 1998, fig. 16). One of the attributes of secondary sexual characteristics is
its appearance during breeding season, followed by regression. Nonetheless, the speci-
mens of A. ojiara examined were collected in May -the end of fall in southern
hemisphere-, and neither males nor females were mature. Therefore, hooks are always
present in the new species; this fact has been observed in the genera Cheirodon and
Hyphessobrycon as well.
In several subfamilies of Characidae such as Iguanodectinae, Stethaprioninae,
Glandulocaudinae or Cheirodontinae and in different lineages within the family, males
usually show hooks on anal and pelvic fins, but sometimes they occur on caudal or
dorsal fins. Within the Cheirodontinae, some species as Serrapinus calliurus bears
hooks in anal fin while many other ones have hooks on anal and pelvic fins. Among
them, C. pisciculus, C. terrabae, C. dialepturus, C. affinis, C. gorgonae (Fink &
Weitzman, 1974), C. ortegai (Vari & Géry, 1980), and C. interruptus (pers. obs.),
Serrapinus kriegi and S. microdon (Uj, 1987; pers. obs.); and Heterocheirodon yatay
A NEW SPECIES OF ASTYANAX FROM ARGENTINA 253
Fics 8-13
Fics 8-10: Astyanax ojiara, lingual view of left teeth, SL 48.0 mm, scale= 0.5 mm. 8: inner row
of premaxillary teeth, with tips gently expanded distally; 9: maxillary tooth with seven small
cusps; 10: eight dentary teeth, decreasing in size anteroposteriorly. Fics 11-13. Astyanax ojiara,
left pectoral axillary scale of males, SL 42.0 mm; 11: lateroventral view, scale= 0.5 mm; 12:
profile of the same scale; 13: inner view, detail of one cleared and stained specimen with 8 hooks,
SL 56.0 mm, scale= 0.5 mm.
254 MARIA DE LAS MERCEDES AZPELICUETA & JOSÉ O. GARCIA
(Casciotta et al., 1992). Hooks are not common in the caudal fin, but, Fink & Weitzman
(1974) described and illustrated hooks not only on the caudal fin of Saccoderma
hastata but also on that fin of Cheirodon dialepturus.
In glandulocaudine species, with notable sexual dimorphism, hooks may occur
on caudal fin, as in Xenurobrycon macropus (Mahnert & Géry, 1984; Weitzman &
Fink, 1985), X. pteropus, Scophaerocharax octopodus, Corynopoma risei, Gephyro-
charax atricaudatus (Weitzman & Fink, 1985), Mimagoniates microlepis and M.
reocharis (Menezes & Weitzman, 1990), Ptychocharax rhyacophila (Weitzman et al.,
1994) Tyttocharax cochui (Weitzman & Ortega, 1995). The presence of hooks in anal
and pelvic fins is usual among glandulocuadine fishes; the position, size and shape of
the hooks vary in different species. Mahnert & Géry (1984) mentioned the occurrence
of hooks on the pectoral fin only in one specimen of X. macropus.
Many other species of characins have been described with hooks; some of them
have hooks on anal fins as Hyphessobrycon diancistrus (Weitzman, 1977a) and H.
procerus (Mahnert & Géry, 1987) or hooks may be present on pelvic fins as in
Paracheirodon axelrodi (Weitzman & Fink, 1983). In most of the species, hooks
develop on anal and pelvic fins: Hyphessobrycon guarani (Mahnert & Géry, 1987), H.
arianne (Uj & Géry, 1989), H. epicharis (Weitzman & Palmer, 1997), H. wajat
(Almirén & Casciotta, 1999), Rachoviscus gracilipes and R. crassipes (Weitzman &
Goncalves da Cruz, 1981), Piabarcus annalis and P. torrenticola, Creagrutus para-
guayensis (Mahnert & Géry, 1988), some Trans-Andean species of Creagrutus as C.
caucanus, C. maracaiboensis, C. paralacus or C. affinis (Harold & Vari, 1994), Hemi-
grammus mahnerti (Uj & Géry, 1989), Bryconamericus theringi (pers. obs.).
Well developed hooks occur also on anal or pelvic fins of other characiform
species such as Brycon microlepis (Gery & Mahnert, 1992), the miniature Priocharax
ariel (Weitzman & Vari, 1987), Toracocharax stellatus (pers. obs.) as well as in some
species of different genera of the Stethaprioninae such as Poptella, Stethaprion,
Brachychalcinus and Orthospinus (Reis, 1989).
In the genus Jupiaba, J. meunieri or J. maroniensis (Gery et al., 1996, as
Astyanax) show hooks on the anal fin. Within Astyanax, hooks on anal and pelvic fins
have been reported in Astyanax maculisquamis (Garutti & Britski, 1997), A. alburnus
(Malabarba, 1983 as A. hasemani), A. kullanderi (Costa, 1995), A. unitaeniatus
(Garutti, 1998), A. eigenmanniorum, A. cf. fasciatus, A. cf. asuncionensis or A. alleni
(pers. obs.). None of those species have caudal fin hooks.
Extremely scarce is the information about the presence of hooks on dorsal fin;
Weitzman (1977b) found hooks on that fin in two species of Hyphessobrycon only, the
Amazonian H. socolofi and H. erythrostigma which bears very few hooks.
The hooks of the pelvic axillary scale appear in medium sized males of A. ojiara
and their number varies in different specimens, although one or two hooks are present
usually (Figs 11, 12). One cleared and stained specimen has eight hooks (Fig. 13) on
that scale. No other characids have been described with hooks on all fins or on pelvic
axillary scale.
A NEW SPECIES OF ASTYANAX FROM ARGENTINA 255
TABLE |
Morphometrics of holotype, 12 females and 12 males (paratypes) of Astyanax ojiara. Minimum,
maximum and mean in parenthesis. DLAR-HJ= distance between last anal-fin ray and hypural
joint.
Holotype females males
Standard length 50.5 39.8-72.0 38.0-53.5
% of standard length
Predorsal distance 51.4 51.8-56.9 (44.8) 50:9-595. 1183)
Preventral distance 49.3 48.7-51.6 (50.4) 47.7-51.3 (48.8)
Preanal distance 64.0 65.8-70.4 (67.7) 61.3-65.9 (63.4)
Body depth 35.0 35.5-40.0 (38.1) 34.0-37.9 (36.5)
Dorsal-fin base 13.4 13.8-17.2 (15.4) 12:8 = 15.9% (1323)
Anal-fin base 32.4 27.6-33.4 (30.5) 30.0-34.6 (32.2)
Pectoral-fin length 23.3 20.3-24.6 (21.7) 22.1-24.5 (23.2)
Pelvic-fin length 19.6 14.8-18.3 (16.5) 17.9-21.5 (19.0)
Distance between pectoral
and pelvic fin origins 2255 20.4-25.7 (23.2) 21.8-24.6 (23.4)
Distance between pelvic
and anal fin origins 19:2 19.6-22.8 (21.3) 16.1-20.0 (18.2)
Head length ZA, 27.3-29.5 (28.1) 26.5-29.7 (27.8)
% of DLAR-HJ
Peduncle depth 90.7 90.0-110.0 (99.0) 74.6-90.7 (83.0)
% of head length
DLAR-HJ 46.4 38.8-46.4 (42.3) 46.8-56.8 (50.2)
Snout length 28.5 24.5-30.9 (27.7) 27.8-30.5 (29.2)
Eye SIN 29.1-37.2 (28.1) 32.3-35.9 (34.3)
Postorbital length SUA 50.0-54.8 (52.4) 53.4-57.2 (55.8)
Interorbital length 32.8 31.0-34.6 (33.0) 31.0-35.6 (32.5)
Maxillary length 22.8 21.1-24.5 (22.8) 37.1-40.6 (38.8)
ACKNOWLEDGMENTS
The authors thank L. R. Malabarba for exchange of specimens of A. eigen-
maniorum, J. R. Casciotta for the photograph, A. Martinez Azpelicueta for the help
with software, J. M. Diaz de Astarloa for making available some literature, and Consejo
Nacional de Investigaciones Cientificas y Técnicas for financial support.
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A NEW SPECIES OF ASTYANAX FROM ARGENTINA 257
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99-136.
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12
REVUE SUISSE DE ZOOLOGIE 107 (2): 259-275; juin 2000
Revision of the genus Ebbrittoniella Martinez
(Coleoptera: Scarabaeoidea: Ceratocanthidae)!
Alberto BALLERIO
Viale Venezia 45, I-25123 Brescia, Italy.
E-mail: ballerio@numerica.it
Revision of the genus Ebbrittoniella Martinez (Coleoptera: Scarabae-
oidea: Ceratocanthidae). - The Oriental genus Ebbrittoniella Martinez,
1962 is redescribed. The current spelling Eubrittoniella is emended.
Cyphopisthes gestroi (Paulian, 1942) is transferred to Ebbrittoniella.
Presently the genus includes E. ignita (Westwood, 1883) and E. gestroi
(Paulian, 1942). Some features of male and female genitalia and the
affinities of the genus are briefly discussed.
Key-words: Coleoptera - Scarabaeoidea - Ceratocanthidae - Ebbrittoniella
- Taxonomy - Oriental Region.
INTRODUCTION
In 1883 Westwood described Acanthocerus (Sphaeromorphus) ignitus on the
basis of a specimen from Sumatra. In the subsequent fifty years Preudhomme de Borre
(1886), Lansberge (1887), Gestro (1899) and Arrow (1912) dealt with this species;
among these authors Gestro for the first time stressed the unnatural generic placement
of the taxon, being the only Oriental representative in a New World genus. In 1962
Martinez, by examining a specimen from Sarawak in BMNH, followed Gestro’s
suggestion and created the new genus Ebbrittoniella to accommodate the taxon ignitus,
while pointing out that the closest genus to it was the Neotropical genus Acanthocerus
Macleay, 1819 (now Ceratocanthus White, 1842).
Paulian in his revision of Oriental Ceratocanthidae (1978) summarized available
distributional data, redescribed the species and placed it in the key near Cyphopisthes
Gestro, 1899. He also emended the name in Eubrittoniella ignitus. This emendation
cannot be considered justified according to the art. 33 of ICZN, because in the original
paper there is no clear evidence of an author’s lapsus calami or printer’s error as
defined by art. 32 of ICZN and therefore the original spelling Ebbrittoniella should be
maintained.
After that point the genus Ebbrittoniella has been listed only by Browne &
Scholtz (1995, 1996, 1998) as Eubrittoniella, in a series of papers upon the hindwing
articulation, base and venation of the Scarabaeoidea.
! Sth contribution to the knowledge of Ceratocanthidae.
Manuscript accepted 28.02.2000
260 ALBERTO BALLERIO
During revisionary studies on Old World Ceratocanthidae I had the opportunity
to examine the type material as well as specimens from several collections, allowing
me to confirm the validity of the genus. Cyphopisthes gestroi (Paulian, 1942) shares all
the generic characters and therefore it is now transferred to Ebbrittoniella.
METHODS AND ACRONYMS
Elytral length is measured from the apex of elytral articular process to the more
external point of the apical convexity, total width measurements are the sum of both
elytral widths.
Drawings of genitalia were taken from pieces previously cleaned with 10%
KOH solution.
Terminology follows Nel & De Villiers (1988) and Nel & Scholtz (1990) for
mouthparts, Kukalovä-Peck & Lawrence (1993) for wing venation and D’Hotman &
Scholtz (1990) for male genitalia (thus the dorsal side of aedeagus is the concave one);
for the remaining conventions I refer to Ballerio (2000).
EE maximum elytral length
EW maximum total elytral width
HL maximum head length
HW maximum head width
È length
PL maximum pronotum length
PW maximum pronotum width
W width
ABCB A. Ballerio private collection, Brescia (Italy)
BMNH The Natural History Museum, London
BPBM Bernice Bishop Museum, Honolulu
HNHM Hungarian Natural History Museum, Budapest
MCSN Museo Civico di Storia Naturale “Giacomo Doria”, Genova
MHNG Muséum d’histoire naturelle, Genève
MNHN Musée national d'Histoire naturelle, Paris
NHMW Naturhistorisches Museum, Wien
NMPC National Museum (Natural History), Praha
OXUM Hope Department of Entomology, Oxford University, Oxford
RMNH Nationaal Natuurhistorisch Museum, Leiden
SACF _ S. Adebratt private collection, Frinnaryd/Boxholm (Sweden)
SMTD Staatliches Museum für Tierkunde, Dresden
ZMUC Zoologisk Museum, Kobenhavns Universitet, Kobenhavn
TAXONOMY
Ebbrittoniella Martinez
Ebbrittoniella Martinez, 1962: 61 (description)
Eubrittoniella Martinez, Paulian, 1978: 506 (emendation)
REVISION OF THE GENUS EBBRITTONIELLA 261
Type species: Acanthocerus (Sphaeromorphus) ignitus Westwood, 1883 by ori-
ginal monotypy.
Etymology: named after Dr. E. B. Britton, former curator at BMNH. The gender
is feminine.
DIAGNOSIS
The genus can be identified by the following combination of characters: labrum
distally abruptly truncate, truncature marked dorsally by a slight carina bearing a row of .
long, erect, fine setae, truncature in frontal view forming a plate irregularly semi-
circular, with semicircumference corresponding to the carina; genal canthus complete,
touching the occipital area; dorsal ocular area large; anterior angles of pronotum
broadly rounded; mesotibiae short and wide (W/L ratio = 0.3), in dd with the inner
apical spur straight and very short; protibiae with outer edge almost smooth (low
magnification), ending with a single tooth in both sexes; apical spur of protibiae sharp
and slender.
DESCRIPTION
Medium to large Ceratocanthidae; volant.
Head: W/L ratio = 1.8, subpentagonal, anterior edge forming a triangle with
obtuse apex (about 150°), both sides of the triangle smooth and almost rectilinear, not
reflexed upward; genae aligned with fore edge, forming a right angle with genal
canthus; genal canthus narrow, straight and complete, touching the occipital area; dor-
sal ocular area large; dorsal interocular area about five times the maximum width of the
dorsal ocular area; ventral ocular area very large; head surface almost plane.
Pronotum:W/L ratio = 1.8, slightly wider than maximum elytral width, evenly
and broadly convex; anterior edge feebly bisinuate; anterior angles distinctly but
slightly protrudent forward, broadly and regularly rounded; sides obtusely rounded;
base narrow, a very thin bead present anteriorly and at each side of base; base with a
weak callosity marking each extremity of base of scutellum.
Scutellum: very large, longer than wide (W/L ratio = 0.7), sides proximally
subparallel and distinctly notched by apical portion of mesepisternum and elytral
articular process, then convergent to form a triangle with apex very elongate and acute
and sides slightly curved inward; apical portion of mesepisterna (see Ballerio, 2000)
visible from above, very large, larger than elytral articular process, subrectangular,
smooth and shiny (Fig. 2d).
Elytra: slightly longer than wide (W/L ratio = 0.9), maximum width near
middle, apex in lateral view fairly reentering inward; slightly flattened on disc, then
abruptly convex at sides to form a pseudoepipleuron not marked by any lateral carina;
marginal elytral area narrow, almost indistinct; elytral suture very feebly raised; sutural
stria very fine and close to suture, limited to distal third; inferior sutural stria absent;
striated articular area well developed and visible in lateral view, relatively wide and
long; elytral articular process well developed, smooth and shiny.
Apical extremity of clypeus (see Ballerio, 2000) short and transversely grooved.
Labrum (Fig. 4b) wide and short, proximally with surface microreticulated, distally
262 ALBERTO BALLERIO
abruptly truncate, truncature marked dorsally by a slight carina bearing a row of long,
erect, distally curled, fine setae; truncature in frontal view forming a plate irregularly
semicircular, with semicircumference corresponding to the carina; surface of plate
almost smooth, bearing few long setae. Distal epipharynx longitudinally divided by a
very sharp strong anterior median process, distally very raised; median brush and
corypha absent; apical fringe made of long fine setae, absent in the middle; lateral
combs made of long fine setae. Mentum ventrally flat, deeply emarginated in the
middle of anterior edge, emargination regularly wide-U-shaped; labial palpi (including
palpiger) four jointed, first joint short and transverse, joint two securiform, joint three
short and ovoidal, the same length of the second, and joint four subconical, about two
times the length of the former, all joints, apart from the last one, fringed with long
setae. Maxillae (Fig. 3a) with a very elongate single lacinia, covered with fine long
setae, monolobed galea proximally sclerotized and distally clothed with very coarse
short thick bi- or triphid setae (galeal brush) (Fig. 3b); maxillary palpi (including
palpiger) four jointed, palpiger very small, joint two wide and relatively short, distinctly
wider than the following joints, joint three about as wide as long, joint four long and
subconical, slightly longer than the preceding two together, apically bearing some short
sensilla. Mandibles elongate, slightly asymmetrical, apicalis more or less gently bent at
about a right angle with apex short and acutely pointed, mesal brush narrow and well
developed, conjunctive present, molar lobe very strong. Antennae 10-jointed, scape
large, distally subcarinate (securiform), distally bearing some setae, funicle short with
pedicellum plump and rounded, the remaining joints very short, distinctly wider than
long, antennal club three-jointed, joints hairy, relatively short, narrow; club small, about
as long as wide and as wide as the length of funicle (L funicle/L lamellae ratio = 2.1).
Ventral areas of prothorax: (Fig. 4a) sides of propleura smooth, very excavated
and folded in, excavation with a further shallow narrow reniform excavation inside
(visible also in dorsal view against the light as a dark reniform patch). Procoxae
transversely oriented, apices nearly touching each other. Anterior trochanters with
anterior tips bearing a tuft of long setae. Profemora slender (W/L ratio = 0.2), posterior
edge without emargination, surface smooth. Protibiae almost straight, outer edge
smooth at low magnfication (at most very slightly serrate: few feeble denticles visible
at 45x); apical spur relatively short, sharp, very gently and feebly curved downward.
Protarsi with first article as long as the following three together, articles two and three
slightly dilated, article five slightly longer than the former; each tarsomere, with the
exception of the last one, ventrally bearing a tuft of dense fine setae. Mesosternum
forming a sharp fine carina, protruding between mesocoxae and joining metasternum.
Mesocoxae almost adjacent to each other, longitudinally oriented. Trochanters with
acute posterior tip. Mesofemora slender (W/L ratio = 0.3), surface smooth, with
posterior edge emarginated at distal third, emargination preceded by a small distinct
tooth. Mesotibiae subrectangular, short (W/L ratio = 0.3), inner angle of apex with two
straight apical spurs. Mesotarsi inserted near the inner angle of apical edge, slightly
longer than apical edge of tibia (exceding it for the length of the last tarsomere), with
first three articles subequal in length, fourth shorter, fifth almost as long as the
preceding two; each tarsomere, with the exception of the last one, ventrally bearing a
263
REVISION OF THE GENUS EBBRITTONIELLA
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264 ALBERTO BALLERIO
tuft of dense setae. Trochanters of metafemora with acute posterior tip, metafemora
plumper than mesofemora (W/L ratio = 0.3), surface wrinkled, posterior edge distally
with a small emargination. Metatibiae triangular, wide (W/L ratio = 0.5), ending with
two straight sharp fine paired spurs. Metatarsi almost as long as the apical edge of tibia,
first article almost as long as the following three together, fifth almost as long as the
first one; each tarsomere, with the exception of the last one, ventrally bearing a tuft of
dense setae.
Wings (Fig. 2a) (Lwing/Lelytron ratio = 2.1): fully developed, MP, ,, - RP loop
present with RP long (although weakly sclerotized), MP, medium sized, apical field
with a vertical secondary sclerification near the radial cell. First axillary with no
appreciable differences at species level (Fig. 2b).
Sexual dimorphism: 2 9 (Fig. 2c: B) have the apical outer tooth of protibiae
distinctly sharper and more protruding outward and forward than in dg d (Fig. 2c: A)
and mesotibiae with the inner apical spur straight and approximately as long as the
outer, while in d d it is straight but very short and very difficult to detect.
Male genitalia: Genital segment (Fig. 5c) Y-shaped (no appreciable differences
at species level), fairly sclerotized, with a distinct manubrium about as long as the basal
triangle, branches forming the manubrium apparently not fused together, although
connected by a transparent membrane (after treatment with KOH), base of triangle very
weakly sclerotized; basal piece of aedeagus large and twisted (Fig. 5a), about four times
the length of parameres; internal sac very large about three times as long as tegmen,
internal sac distally with coarse spicules and setae; temones present; parameres (Fig.
5b) short and slightly asymmetrical (hardly appreciable differences at species level),
laterally flattened, between parameres dorsally lies a narrow subtriangular sclerite.
Female genitalia: bursa copulatrix (Fig. 5d) with one small symmetrical sub-
circular or subtriangular sclerite (very variable in shape and without appreciable
differences at species level) with a hole in the middle; spermatheca (Fig. 6) strongly
sclerotized, large and distinctly wide-U-shaped; genital palpi weakly sclerotized, sub-
circular, relatively small and short.
Ebbrittoniella ignita (Westwood, 1883) Fig. la
Acanthocerus (Sphaeromorphus) ignitus Westwood, 1883: 2 (description)
Acanthocerus (Sphaeromorphus) ignitus Westwood: Preudhomme de Borre, 1886: 80 (catalogue)
Synarmostes ignitus (Westwood): Lansberge, 1887: 209 (list)
Acanthocerus ignitus Westwood: Gestro, 1899: 462 (redescription, distribution and key)
Acanthocerus ignitus Westwood: Arrow, 1912: 49 (catalogue)
Ebbrittoniella ignita (Westwood): Martinez, 1962: 61 (redescription and iconography)
Eubrittoniella ignitus (Westwood): Paulian, 1978: 506 (redescription, distribution, iconography
and key)
Type locality: Koetoer (Indonesia: Sumatra).
Material examined - Holotypus d: Koetoer 6.78 / Sum. Exp. Koetoer 6.78 / Typus /
Acanthocerus ignitus Westw. / Museum Leiden Acanthocerus ignitus det. Westwood / 6903 /
Eubrittoniella ignitus (Westw.) R. Paulian det. / Ebbrittoniella ignita (Westw.) A. Ballerio det.
1997 (RMNH), [very damaged specimen, pinned, lacking the head, which is glued together with
legs and mouthparts on a card pinned with a separate pin, without any data bearing label].
REVISION OF THE GENUS EBBRITTONIELLA 265
b Coi
A
Fic. 2 — a: E. gestroi (Cameron Highlands), wing: vertical secondary sclerification (S) (scale bar:
1 mm); b: E. gestroi (Cameron Highlands), first axillary (scale bar: 0,1 mm; c: Ebbrittoniella,
apex of foretibiae (dorsal view): (A) male, (B) female; d: Ebbrittoniella, area near scutellum as
seen when the beetle assumes the “rolled up” posture: apical portion of mesepistemum (M),
articular process of elytron (AP), scutellum (S), elytron (E), pronotum (P).
Other 32 specimens (7 dd and 4 9 2 dissected). INDONESIA. Kajoe Tanam [Sumatra],
Klein (ZMUC); Sumatra, Manna, M. Knapper (RMNH); Ruuyer Payakor, Sumatra (RMNH);
Sentinjak, Sumatra, 1800 ft., Jan. ‘98, 99-95 (BMNH); Setinjak, W. Sumatra, I to VII. 98,
(Ericsson) (Coll. C. Felsche, Kauf 20, 1918) (SMTD); Sumatra, Padang Sidempoean, XII.1902-
1.1903 (MNHM); Sumatra (ex Mus. Van Lansberge) (MNHM). Nias Island. Is. Nias, U. Raap
(MNHM); Hili Madjedja, N. Nias, 4me trim. 195, I.Z. Kannegieter (ex Mus. Van de Poll)
(MNHM); Is. Nias 1897-98, U. Raap (RMNH); Is. Nias 1897-98 U. Raap (MCSN). Kalimantan-
Timur: Apokayan, Long Sungei Barang 900 m, secondary forest, 15-23.02.1997, leg. C. & P.
Zorn. MALAYSIA. Malay penin.: Selangor, Giunting Simpak, Jan. 2"¢ 1933, N.M. Pendlebury,
266 ALBERTO BALLERIO
F.M.S. Museum (BMNH); Malaysia: Pahang/Johor, Endau Rompin NP, 100 m, Salendang, 28.2-
12.3 1995, leg. Strba & Hergovits (NHMW); Malaysia — Pahang, Banjaran Benom, Lata Jarom,
6-8.3.1997, Ivo Jenis leg. (ABCB); ibidem, 18-21.3.1997, Ivo Jenis leg. (ABCB), W. Malaysia,
Pahang, Baniaran Benom Mts., K. Ulu Dong 10-15 km SSE, 17-23.1V.1997, D. Hauck leg.
(ABCB); Sarawak, C. S. Brooks, B.M. 1928-193 (BMNH); Malaysia: Sabah, Sipitang, Mendo-
long, T1B/W4, 11.V.1988, leg. S. Adebratt, 12-60, 2859BC (SACF). OTHER. Sunda-Inseln v.
Studt /coll. Petrovitz (MHNG).
DESCRIPTION
HL= 0.8-1.2 mm HW= 1.6-2 mm PL= 1.5-2 mm PW=2.8-3.3 mm EL= 2.9-3.9 mm
EW= 2.7-3.6 mm.
Dorsally very shiny and entirely brightly metallic: red with gold/green faint,
elytral suture, scutellum, sides of elytra, sides of pronotum and sides of head deep
electric blue with green faint; ventrally alutaceous, reddish brown, antennae yellowish.
Pubescence invisible at low magnification, otherwise formed by very short and fine
hyaline erect setae, subject to wear. Head surface with a variable number of transverse
lines, occupying anteriorly the two thirds of head; vertex with sparse simple punctures,
often preceded by a very short transversal impressed line. Pronotum very convex,
surface with very small horseshoe-shaped punctures (containing a small simple
puncture in the middle) or sometimes simple punctures, never very dense, usually
sparser on disc and slightly coarser near anterior angles; the horseshoe is opened
outwards in the punctures at sides and inwards in the punctures on the disc. Scutellum
with horseshoe-shaped punctures with horseshoe opened toward apex, punctures denser
near sides. Elytra distinctly convex, surface with punctures horseshoe-shaped, small,
never very dense, variably distributed, horseshoe opened toward suture or apex; two
longitudinal lines along the suture at medial and distal third of length; pseudoepipleura
with some longitudinal lines; humeral callus not protruding; articular area very large.
Outer face of meso- and metatibiae with short impressed lines varying from longitu-
dinal to transverse and short erect setae through the entire surface. Wings with vertical
secondary sclerification of apical field finer than in E. gestroi and MP,,, - RP loop
shorter. Spermatheca as in figs. 6a, b, c, d.
VARIABILITY
Although the majority of individuals examined shows the typical colours, one
pair from Pahang is uniformly dark green, one individual from Nias is uniformly deep
amaranth and the individual from Sabah is yellowish with greenish metallic sheen;
there is a strong variability also in the punctures (mostly the ones of pronotum), which
vary in size and density: the specimen from Sabah has the entire head surface covered
by transverse lines, the amaranth specimen from Nias has the entire pronotum
completely smooth, while the pair of dark green specimens has punctures coarser and
more impressed than the other specimens. The very fine pubescence varies and in
some specimens is no longer visible (probably ripped off by wear).
DISTRIBUTION AND HABITAT
Recorded from Peninsular Malaysia, Sumatra, Nias Island and Borneo. For
habitat see under E. gestroi.
REVISION OF THE GENUS EBBRITTONIELLA 267
Fic. 3 — a: E. gestroi (Cameron Highlands), maxilla (SEM micrograph); b: E. gestroi (Cameron
Highlands), detail of galeal brush (SEM micrograph).
268 ALBERTO BALLERIO
REMARKS
°°
Westwood (1883) described also a variety “paullo minor, obscurior,...” from
Sarawak without naming it; examination of the specimen in OXUM revealed that it is
an Eusphaeropeltis sp.
Ebbrittoniella gestroi (Paulian, 1942) comb. n. Fig. 1b
Philharmostes Gestroi Paulian, 1942: 70 (description and key)
Cyphopisthes gestroi (Paulian): Paulian, 1978: 512 (new combination, redescription, distribution,
iconography and key)
Type locality: Palembang (Indonesia: Sumatra).
Etymology: named after Dr. R. Gestro, late director of MCSN and specialist of
Ceratocanthidae.
Material examined — Holotypus, 2: Sumatra Palembang / Type / Cyphopisthes gestroi n.
sp. det. R. Paulian, 1937/ Ebbrittoniella gestroi (Paulian) n. comb. det. A. Ballerio, 1997.
(MNHN) [completely rolled up and glued on a card] .
Other 141 specimens ( 15 dd and 15 9 9 dissected) - INDONESIA. Palembang
[Sumatra], 1900, Bouchard (MCSN); MALAYSIA. Malaya: (W) Perak, Maxwell Hill, 1350 m.,
17-20.1IL.1958, T.C.Maa collector, Bishop (BPBM); Malaysia — Perak, Banjaran Bintang,
Maxwell Hill (Taiping), 18-19.2.1997, leg. Ivo Jenis (ABCB); West Malaysia, Perak, Maxwell
Hill, 900-1000 m, above Taiping City, 12-16.1.1995, leg. S. Beévar j. & s. (ABCB); W.
Malaysia, Perak: 25 km NE of Ipoh, 2100 m, Banjaran Titi Wangsa mts., Gunung Korbu, 4-
13.111.1998, P. Cechovskÿ leg.; W. Malaysia, Perak, 25 km NE of Ipoh, 1200 m, Banjaran Titi
Wangsa mts., Gunung Korbu, 27.1-2.11.1999, P. Cechovsky leg.; W. Malaysia, Pahang, 30 km E
of Ipoh, 1500 m, Cameron Highlands, Tanah Rata, 20.11-3.1II. 1998, P. Cechovsky leg.; ibidem,
22-26.1.1999, P. Cechovsky leg.; P. Malaysia-Pahang: Cameron Highlands, Tanah Rata umg.,
gn. Jasar, 1300 m, 25.11.1997, leg. Schuh & Lang (ABCB); West Malaysia, Pahang, C.
Highlands, Tanah rata, 20-25.1.1995, Gn. Jasar, 14-1500 m, leg. S. Beëvar j. & s. (ABCB); West
Malaysia, Pahang, Cameron Highlands, Brinchang, 18-19.1.1995, Gunung Berembam, 1600 m,
leg. S. Beévar j. & s. (ABCB); Malaysia, Pahang, Cameron Highlands, 2 km S of Tanah Rata on
Tapah Road, montane rainforest, at light no. 93, 29.11.1995, leg. O. Merkl & L. Szikossy
(HNHM); Malaysia, Pahang, Cameron Highlands, Tanah Rata, from illuminated white washed
walls no. 77, 23-31.111.1995, 1. O. Merkl (HNHM); Malaysia — Pahang, Banjaram Benom, Lata
Jarom, 6-8.3.1997, leg. Ivo Jeni$ (ABCB); Malaysia — Perak, Cameron Highlands, Tanah Rata,
13-16.3.1997, leg. Ivo Jenis (ABCB); Borneo (Brit. N.), Sandakan bay (SW), Sapagaya Lumber
Camp, 2-20 m, XI-3-157, J.L. Gressit Collector (BPBM); North Borneo (SE), Forest Camp, .19
km N. of Kalabakan, 18.X1.1962, Y. Hirashima, Light trap, Bishop (BPBM); ibidem, 7-
10.X1.1962 (BPBM); ibidem, 27.X.1962 (BPBM): ibidem, 10.X.1962 (BPBM); Malaysia —
Sabah, Crocker Range National park, Longkogungan env., Ca. 750-850 m a.s.l., 19-21.VI.1996,
7c (NHMW); Borneo: Sarawak Bau District, Bidi, 90-240 m. 3.IX.1958, T.C. Maa Collector
Bishop (BPBM); Sarawak, Kapit dist., Sebong, Baleh riv., 9-21.3.1994, Sv. Bily leg. (NMPC),
Sarawak, Kapit dist., Rumah Ugap vill., Sut. Riv., 3-9.3.1994, Sv. Bily leg. (NMPC); Sarawak:
Gunung Mulu Nat. Park, R.G.S. Exped. 1977-8, J.D. Holloway et al. B.M. 1978-206, site 2,
january, camp 4, Mulu, 1790 m, 452463, lower montane (moss) forest, acl-understorey (BMNH).
DESCRIPTION:
HL= 0.9-1.1 mm HW= 1.5-1.9 mm PL= 1.3-1.8 mm PW=2.7-3.2 mm EL= 3-3.5 mm
EME 232 mms:
Dorsally metallic bronze/green with reddish faint, head, pronotum, scutellum
and elytra covered by long recumbent yellowish setae; ventrally alutaceous yellowish/
brown with antennae yellowish. Head as in E. ignita but completely covered by
recumbent setae and dense punctures. Pronotum slightly less convex than in E. ignita,
REVISION OF THE GENUS EBBRITTONIELLA 269
FIG. 4 — a: E. gestroi (Cameron Highlands): ventral areas of prothorax (SEM micrograph); b: E.
gestroi (Cameron Highlands): head in fronto-lateral view, showing the truncature of labrum
(SEM micrograph).
270 ALBERTO BALLERIO
surface completely covered by very dense small horseshoe-shaped punctures, with
horseshoe very short and almost transversal; each horseshoe with a small impressed
puncture in the middle; pubescence very long and dense, recumbent, some subcircular
small and weak depressions at each side of disc. Scutellum with coarse horseshoe-
shaped punctures and long recumbent pubescence. Elytra slightly less convex than in
E. ignita, surface with punctures less dense and slightly larger than on pronotum,
relatively sparser near suture; proximal third with large longitudinal smooth areas
between the suture and the pseudoepipleura; pseudoepipleura with transversal long
dense horseshoe-shaped punctures. Humeral callus very developed and protuding
outwards; articular area smaller than in FE. ignita. Outer face of meso- and metatibiae
with short impressed lines varying from longitudinal to transverse and short erect setae
through the entire surface. Wings (Fig. 2a): vertical secondary sclerification of apical
field thicker than in E. ignita and MP, ,, - RP loop distinctly longer. Spermatheca as in
figs. 6e, f, g, h.
VARIABILITY
E. gestroi is less variable compared to the former species; the colours vary from
bronze green to green with faint reddish sheen. In the pair from Lata Jarom the smooth
elytral areas are very small, while punctures and lines are slightly more impressed.
DISTRIBUTION AND HABITAT
Recorded from Peninsular Malaysia, Sumatra and Borneo (Sabah and Sarawak).
Both species are found in rainforests, they are sympatric and sometimes also
syntopic (Lata Jarom, leg. Jeni$). E. ignita seems to be restricted to lowland rainforests,
where it has been collected by beating leaves or with window traps; E. gestroi shows a
broader ecological range because it is found in both lowland and montane forests (till
2100 m a.s.l.), where it is the most commonly collected Ceratocanthidae. There are
several records of this species at light. The unusual vestiture of galeal brush, which is
covered by short and thick bi- or triphids dense setae (while in most other Cerato-
canthidae these setae are longer, finer and usually sharp) could suggest that Ebbritto-
niella has feeding habits different from the ones of the majority of Ceratocanthidae.
KEY TO THE SPECIES OF EBBRITTONIELLA
I Labrum distally distinctly truncate, truncature marked dorsally by a
slight carina bearing a row of long, fine, erect setae, truncature in frontal
view forming a plate irregularly elliptical or semicircular................. 2
- Labrum without distinct truncature marked dorsally by a carina
DD RE ease uae 20 UNS NEE RISE ERE NEN INA i other Ceratocanthidae
2 Anterior angles of pronotum triangular; mesotibiae slender and relatively
narrow (W/L ratio = 0,2), with inner apical spur of d d bent inwards at a
right angle; apex of protibiae usually ending with two teeth in the 9 9
(one known exception); apical spur of protibiae strong, apically distinctly
Pent downwards ohio) Met ee ee ee eee Cyphopisthes Gestro
REVISION OF THE GENUS EBBRITTONIELLA 271
- Anterior angles of pronotum broadly rounded; mesotibiae short and wide
(W/L ratio = 0,3), with the inner apical spur of d d straight and extre-
mely short; apex of protibiae ending with a single tooth in both sexes;
apical spur of protibiae fine, very gently and regularly bent downwards.
TT SE Ce me Sy tae ao Ebbrittoniella Martinez
3 Surface of pronotum strongly punctured, punctures very dense, giving a
granulose appearance, pronotal pubescence long, yellowish, recumbent,
elytra with the same microsculpture and pubescence as pronotum, but
with some smooth longitudinal areas, humeral callus very pronounced
WET CRIE E. gestroi (Paulian)
- Surface of pronotum smooth, with sparse simple or horseshoe shaped
punctures, glabrous or at most with very fine and short erect hyaline
pubescence, elytra like pronotum, humeral callus weakly pronounced
De D MR SR RS I AE NER: E. ignita (Westwood)
DISCUSSION
Some interesting new discoveries about the genitalia of the Ceratocanthidae
were made during this revision. First of all it was found that the bursa copulatrix, which
contains the spermatophore, bears a small subcircular sclerite on its inner wall, which
probably has the function of breaking the spermatophore; similar sclerites have been
observed in other genera of Ceratocanthidae, namely Pterorthochaetes Gestro, 1899
(Ballerio, 1999), Philharmostes Kolbe, 1895 and its allies (Ballerio, 2000) and
Eusphaeropeltis Gestro, 1899 (author’s unpublished data). Secondly a mobile acces-
sory sclerite was detected, lying between the ventral basal extremities of parameres and
associated with the median lobe; normally it lies parallel to the anchoring point of
parameres with basal piece, but, when the internal sac is everted, the sclerite is raised at
right angle, perpendicular to the anchoring point of parameres; it is possibly an
intermediate form between an anchoring and a supporting sclerite (see D’Hotman &
Scholtz, 1990).
In order to find the closest relationships of the genus Ebbrittoniella an analysis
was conducted on all the genera of typical Ceratocanthidae, on the basis of all available
morphological characters of adults, using the sister group of Ceratocanthidae, i.e. the
Hybosoridae (Browne & Scholtz, 1996) and in particular the genera Phaechrous
Castelnau, 1840 and Liparochrus Erichson, 1848 (Howden & Gill, 1995), as an out
group for character polarization. The analysis suggests that Cyphopisthes Gestro, 1899
could be the genus closest to Ebbrittoniella. The following synapomorphies are shared
by the two genera: a) outer edge of protibiae smooth (low magnification), b) sexual
dimorphism involving the apical teeth of protibiae, c) labrum distally truncate, trun-
cature marked dorsally by a slight carina bearing a row of erect fine setae, d) sperma-
theca strongly sclerotized and e) metathoracic wings: apical field with a vertical secon-
dary sclerification near the radial cell. The genus Cyphopisthes can be divided into two
groups, whose definition and status are being dealt with in a separate paper, Ebbritto-
niella shares four further synapomorphies with the group formed by C. acromialis
272 ALBERTO BALLERIO
Fic. 5 — a: E. gestroi (Cameron Highlands), aedeagus with accessory sclerite (AS) (scale bar: 1
mm); b: E. gestroi (Cameron Highlands), parameres (lateral view) (scale bar: 0,1 mm); c:
E. gestroi (Cameron Highlands) genital segment (scale bar: 1 mm); d: E. gestroi (Cameron
Highlands), bursa copulatrix with sclerite (S) and spermatophore inside (scale bar: 1 mm).
REVISION OF THE GENUS EBBRITTONIELLA 273
h
Fic. 6 — a (Lata Jarom), b (Lata Jarom), c (Lata Jarom), d (Sabah: Mendolong): E. ignita,
spermatheca; e (Borneo: north of Kalabakan), f (Maxwell Hill), g (Cameron Highlands), h
(Cameron Highlands): E. gestroi, spermatheca (scale bar: 0,1 mm).
274 ALBERTO BALLERIO
(Pascoe, 1860) and few other species, i.e. f) labium with a very deep and wide regular-
U-shaped excavation in the middle, g) spermatheca deeply U-shaped, h) bursa
copulatrix with a characteristically shaped sclerite, and i) presence of an accessory
sclerite among parameres. Characters b), c) and h) are autoapomorphic.
The similarities with Ceratocanthus White, 1842 stressed by earlier authors are
likely due to convergent evolution and, although at the present stage of knowledge
remote relationships cannot be excluded, there are several characters that place
Ebbrittoniella and Cyphopisthes quite far from Ceratocanthus. The latter differs at least
in the following characters: a) labrum neither abruptly truncate nor divided by any
carina, b) elytra without a false epipleuron, c) metathoracic wings: MP,,, — RP loop
absent, d) metathoracic wings: apical field without vertical secondary sclerification, e)
meso- and meta- tarsi capable of being folded along the longitudinal axis of the inner
face of the tibia, f) sexual dimorphism not involving the shape of apical teeth of
protibiae and g) spermatheca weakly sclerotized.
ACKNOWLEDGEMENTS
I wish to thank the following people for permission to study material, loan of
specimens, valuable information and advice: S. Adebratt (Frinnaryd, Sweden), D.
Ahrens (SMTD), S. Beévai (Ceské Budéjovice, Czech Republic), S. Bily (NMPC), Y.
Cambefort (MNHN), P. Cechovsky (Brno, Czech Republic), G. Cuccodoro (MHNG), I.
Jenis (Naklö, Czech Republic), M.D. Kerley (BMNH), J. Krikken (RMNH), I. Löbl
(MHNG), O. Martin (ZMUC), G. McGavin (OXUM), O. Merkl (HNHM), O.
Montreuil (MNHN), R. Paulian (Bordeaux), R. Poggi (MCSN), G. A. Samuelson
(BPBM), C.H. Scholtz (University of Pretoria), J. Scheuern (Westum, Germany), H.
Schönmann (NHMW), R. Schuh (Katzelsdorf, Austria), C. Zorn (Dresden). R. Furlan
(Centro C.U.G.A.S., Universita di Padova) took SEM micrograph and J. Kobylak
(Praha) executed the habitus drawings.
REFERENCES
Arrow, G. J. 1912. Coleopterorum Catalogus pars 43 - Scarabaeidae: Pachypodinae, Pleoco-
minae, Aclopinae, Glaphyrinae, Ochodaeinae, Orphninae, Idiostominae, Hybosorinae,
Dynamopinae, Acanthocerinae, Troginae. W. Junk, Berlin, 66 pp.
BALLERIO, A. 1999. Revision of the genus Pterorthochaetes, first contribution (Coleoptera:
Scarabaeoidea: Ceratocanthidae). Folia Heyrovskyana 7(5): 221-228.
BALLERIO, A. 2000. A new genus and species of Ceratocanthidae from Tanzania (Coleoptera:
Scarabaeoidea). African Zoology 35(1): 131-137.
BROWNE, D. J. & SCHOLTZ, C. H. 1995. Phylogeny of the families of Scarabaeoidea (Coleoptera)
based on characters of the hindwing articulation, hindwing base and wing venation.
Systematic Entomology 20: 145-173.
BROWNE, D. J. & SCHOLTZ, C. H. 1996. The morphology of the hind wing articulation and wing
base of the Scarabaeoidea (Coleoptera) with some phylogenetic implications. Bonner
zoologische Monographien 40: 1-200.
BROWNE, J. & SCHOLTZ, C. H. 1998. Evolution of the scarab hindwing articulation and wing base:
a contribution toward the phylogeny of the Scarabaeidae (Scarabaeoidea: Coleoptera).
Systematic Entomology 23: 307-326.
REVISION OF THE GENUS EBBRITTONIELLA 275
D’HoTMan, D. & SCHOLTZ, C. H. 1990. Phylogenetic significance of the structure of the external
male genitalia in the Scarabaeoidea (Coleoptera). Entomology Memoir Department of
Agricultural Development Repubublic of South Africa 77,51 pp.
GESTRO, R. 1899. Sopra alcune forme di Acanthocerini. Annali del Museo civico di Storia natu-
rale di Genova XXXIX: 450-498.
HOWDEN, H. F. & GILL, B. D. 1995. Trachycrusus, a new genus of Ceratocanthinae (Coleoptera
Scarabaeidae) with two new species from Peru. The Canadian Entomologist 127:
587-593.
KUKALOVA-PECK, J. & LAWRENCE, J. F. 1993. Evolution of the hind wing in Coleoptera. The
Canadian Entomologist 125: 181-258.
LANSBERGE, J. W. VAN 1887. Trogides nouveaux. Notes from the Leyden Museum 1X:199-211.
MARTINEZ, A. 1962. Un nuevo género de Acanthocerinae (Col. Scarabaeidae). Physis 23 n. 64:
61-64.
NEL, A. & DE VILLIERS, W. M. 1988. Mouthpart structure in Adult Scarab Beetles (Coleoptera
Scarabaeoidea). Entomologia Generalis 13 (1/2):95-144.
NEL, A. & SCHOLTZ, C. H. 1990. Comparative morphology of the mouthparts of adult Scara-
baeoidea (Coleoptera). Entomology Memoir Department of Agricultural Development
Repubublic of South Africa 80, 84 pp.
PAULIAN, R. 1942. Coléoptéres Acanthocérides nouveaux ou peu connus. Revue francaise
d’Entomologie IX: 70-75.
PAULIAN, R. 1978. Revision des Ceratocanthidae [Col. Scarabaeoidea] II - Les espèces orientales
et australiennes. Annales de la Societe entomologique de France (N.S.) 14 (3): 479-514.
PREUDHOMME DE BORRE, A. 1886. Catalogue des Trogides décrits jusqu’à ce jour, précédé d’un
synopsis de leur genres et d’une esquisse de leur distribution géographique. Annales de la
Société entomologique de Belgique 30: 54-82.
WESTWOOD, J. O. 1883. Two new species of the coleopterous genus Acanthocerus. Notes from
the Leyden Museum 5: 1-2.
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REVUE SUISSE DE ZOOLOGIE 107 (2): 277-281; juin 2000
A new desert psocid from Namibia (Insecta: Psocoptera: Trogiidae)
Charles LIENHARD
Museum d’histoire naturelle, c. p. 6434, CH-1211 Geneve 6, Switzerland.
E-mail: charles.lienhard @ mhn.ville-ge.ch
A new desert psocid from Namibia (Insecta: Psocoptera: Trogiidae). -
A single female of a new species representing a new genus of the family
Trogiidae (Psocoptera: Trogiomorpha: Atropetae) is described and illus-
trated: Spinatropos philippi gen. n., sp. n. The specimen has been collected
in an unusual biotope for Psocoptera, at Diaz Point (Liideritz), on a rocky
desert headland with hardly any vegetation.
Key-words: Psocoptera - Trogiidae - new genus - new species - desert
fauna - Namibia.
INTRODUCTION
Almost nothing is known on Psocoptera of Namibia. The only psocid recorded
from this country (from Windhoek) is the cosmopolitan and usually domestic species
Liposcelis bostrychophila Badonnel, 1931 (cf. Checklist of Southern African Insects on
Internet: http://www.ru.ac.za/departments/zooento/Martin/Insects.html).
The present material has been collected by N. P. & M. J. Ashmole under stones
on a rocky desert headland with hardly any vegetation near Diaz Point (Liideritz), a
very unusual biotope for Psocoptera. These insects generally live on vegetation, mostly
trees or shrubs, or in leaf-litter (cf. Lienhard, 1998a). An important factor which enables
a psocid to live in the above mentioned biotope could be the proximity to the sea, which
guarantees a constantly high relative humidity of the air. The occurrence of a psocid of
the genus Liposcelis Motschulsky, 1852 on apparently “sterile” littoral limestone rocks
in the Mediterranean (Cyprus) has been reported by Lienhard (1998b). It would be
worthwhile for ecologists to pay more attention to psocids accidentally captured in such
unusual situations.
Besides these ecological aspects the present material is also interesting from a
taxonomic and perhaps ethological point of view. It represents a very characteristic new
species of the family Trogiidae which cannot be placed in one of the hitherto known
genera of this family. Its most striking diagnostic character, the presence of two short
stout externoapical spurs on each of the somewhat elongated third valvulae (fig. 1) may
prove to be connected with special ovipositional behaviour. As known up to now, all
Psocoptera lay their eggs on the surface of a substrate (cf. Lienhard, 1998a). The new
species, however, seems to be equipped to slightly dig in its eggs into a sandy soil.
Manuscript accepted 18.01.2000
278 CHARLES LIENHARD
According to N. P. Ashmole (in litt.) there were small deposits of sand or fine rock
fragments in the crevices of its habitat at Diaz Point. Similar digging spines on
ovipositor are known in other insects, e.g. Diptera, where they are present in sand-
dwelling Therevidae (cf. Irwin & Lyneborg, 1981: p. 514, fig. 22) or in the essentially
dune-inhabiting species Helina protuberans Zett. (Muscidae), which lays its eggs into
sandy soils (cf. Hennig, 1964: p. 209, Textfig. 46).
Spinatropos gen. n.
Diagnosis. Apterous, mesothorax in dorsal view about of same width as
prothorax, metathorax distinctly wider. Body densely pubescent, excepting the almost
bald postclypeus, which bears only a few peripheral hairs. Postclypeus of normal shape,
not exceedingly bulging. Compound eyes well developed, hemispherical, with some
small hairs in dorsal half, ocelli absent. Maxillary palpus (fig. 4) with fourth (terminal)
segment (P4) of cylindrical shape, some stout setae present in apical half of P2, P3 and
P4, no forked sensillum on P4. Laciniae symmetrically developed, lacinial tip
tridentate, teeth diverging and well developed (fig. 3). Hind tibia with two apical spurs
and two spurs on inner face in apical half. Pretarsal claw lacking preapical tooth,
pulvillus long and slender, its tip slightly enlarged (fig. 6). 8th sternite of the female
without sclerified knob which could be used in sound production (cf. Lienhard, 1998a:
fig. 26m, Trogium pulsatorium). Third valvula (fig. 1) somewhat elongated, as typical
of the family-group Atropetae, with spiny pilosity, two short stout externoapical spurs
and one slender normal seta near tip. Second valvula (fig. 1) relatively well developed,
reaching almost to the middle of the third valvula. Spermapore simple (fig. 5).
Spermatheca with two oval parietal glands of about equal dimensions, situated close to
each other in the proximal zone of the vesicle (fig. 7). Glands with numerous pores, no
papillae (fig. 8). Spermatophore with very long tubular part (fig. 7). Male unknown.
Type species. S. philippi sp. n.
Etymology. Spina (lat.) = spine, an allusion to the particular pilosity of the third
valvula. Atropos = one of the three Fates, the antique goddesses of destiny; Atropos
Leach, 1815 is a synonym of Trogium Illiger, 1798 and gave rise to the family-group
name Atropetae. In analogy to Atropos the new name is of feminine gender.
Discussion. See discussion of the type species.
Spinatropos philippi sp. n. Figs 1-8
MATERIAL
Holotype 9. Namibia: Diaz Point (Liideritz), rocky desert biotope with hardly
any vegetation, relatively close to the sea, by visual seaching under stones, 30.X1.1994,
leg. N. P. & M. J. Ashmole (sample nr 3090) (Muséum d'histoire naturelle de Genève).
ETYMOLOGY
The species is dedicated to N. Philip Ashmole, one of its collectors, in
acknowledgement of his tireless efforts in collecting psocids in biotopes where they
“should not occur”.
A NEW DESERT PSOCID FROM NAMIBIA 279
DESCRIPTION (2)
Coloration. Body, legs and antenna whitish to light brown. Compound eye black
(after 4 years in alcohol). Apical half of flagellar segments darker brown than basal half
in basal part of the antenna, flagellar segments in its apical part entirely brown.
Morphology. See generic diagnosis, with the following complements. Vertical
and frontal sutures distinct. Both antennae damaged, one with 22, the other with 11
segments. Marginal sensilla of labrum (fig. 2) typical of the family Trogiidae (cf.
Badonnel, 1977). The complete aptery of the specimen is real and not due to broken
winglets. This is supported by the absence of insertion points of winglets and by the
even pilosity of thoracic tergites (in the frequently encountered cases of micropterous or
brachypterous Trogiidae where winglets are broken, insertion points are visible on
mesothoracic tergite and the lateral parts of the metathoracic tergite are bald, as it has
been figured for the genus Cerobasis Kolbe, 1882 by Lienhard, 1998a: fig. 24c).
Mesothorax (in dorsal view) shorter than half width of vertex. No coxal organ differen-
tiated. Femora somewhat thickened, hind femur about three times as wide as hind tibia.
Hind tibia with some long hairs on outer face. First segment of hind tarsus with
numerous stout spur-like setae on ventral face. Epiproct bald in the middle, with some
lateral hairs. Paraprocts with a distinct anal spine on hind margin. Subgenital plate
absent, ovipositor valvulae basally covered by some membraneous folds. The wall of
the spermathecal vesicle bears numerous fine pores, especially in the more distal half
(fig. 7). The length of the oval spermathecal glands is about 125 um, each gland bears
much more than 100 pores (fig. 8).
Measurements. Body length = 2,0 mm. Vertex (width of head capsule) = 650
um. Length of hind femur = 520 um. Length of hind tibia = 750 um. Length of hind
tarsomeres (measured from condyle to condyle): tl = 320; t2 = 72; t3 = 90.
Remark. The male is not yet known, but the presence of a spermatophore in the
spermatheca of the female gives evidence of the bisexuality of the species.
DISCUSSION
Within the family Trogiidae (sensu Smithers, 1990) the new genus is charac-
terized by the very particular spiny pilosity of the third valvula, especially by the
presence of two stout externoapical spurs, but also by the cylindrical shape of the
terminal segment (P4) of the maxillary palpus. In all other non-fossil genera, P4 is at
least slightly enlarged apically, often much enlarged (hatchet-shaped). In the genera
where females are known (not known in Anomocopeus Badonnel, 1967) the pilosity of
the third valvula consists of normally shaped slender setae, as usual in the family-group
Atropetae. The genus can also be distinguished from most of the other genera (amber
fossils included) by its complete aptery. The only other apterous genera are Anomo-
copeus and Mymicodipnella Enderlein, 1909; some apterous species are also known in
the large genus Cerobasis.
Nothing exact is known about the biology of the new species, for some general
remarks, see the introduction.
280 CHARLES LIENHARD
Fics 1-8
Spinatropos philippi gen. n., sp. n., female: 1, ovipositor valvulae; 2, marginal sensilla of labrum;
3, lacinial tip: 4, maxillary palpus: 5, spermapore; 6, pretarsal claw: 7, spermatheca, containing
one spermatophore: 8, parietal gland of spermatheca.
A NEW DESERT PSOCID FROM NAMIBIA 281
ACKNOWLEDGEMENTS
I am very grateful to N. Philip and Myrtle J. Ashmole (Edinburgh) for having
made their Namibian collections of psocids available to me. I also thank N. P. Ashmole
for reading the manuscript and making some valuable suggestions. I am indebted to my
colleague Bernhard Merz (Geneva) for information about the presence of digging
spines on the ovipositor in Diptera.
REFERENCES
BADONNEL, A. 1977. Sur les sensilles du bord distal du labre des Psocoptères et leur intérêt
taxonomique. Bulletin de la Société entomologique de France 82(5-6): 105-113.
HENNIG, W. 1964. Muscidae (pp. 1-624). In: LINDNER, E. (ed.). Die Fliegen der palaearktischen
Region. Vol. 7(2), part 63b(1).
IRWIN, M. E. & LYNEBORG, L. 1981. Therevidae (pp. 513-523). In: Manual of Nearctic Diptera.
Vol. 1. Research Branch Agriculture Canada, Monograph 27: VI + 674 pp.
LIENHARD, C. 1998a. Psocopteres euro-méditerranéens. Faune de France 83: XX + 517 pp.
LIENHARD, C. 1998b. Interessantes aus der Welt der Psocopteren. Mitteilungen der Schwei-
zerischen Entomologischen Gesellschaft 71(1-2): 252-253.
SMITHERS, C. N. 1990. Keys to the families and genera of Psocoptera (Arthropoda: Insecta).
Technical Reports of the Australian Museum 2: 1-82.
Eu
L
REVUE SUISSE DE ZOOLOGIE 107 (2): 283-323; juin 2000
Terrestrial Isopoda from Guatemala and Mexico
(Crustacea: Oniscidea: Crinocheta)
Andreas LEISTIKOW
Universität Bielefeld
Abteilung für Zoomorphologie und Systematik
Morgenbreede 45
D-33615 Bielefeld.
E-mail: Leiste@Biologie.Uni-Bielefeld.de
Ruhr-Universität Bochum
Lehrstuhl für spezielle Zoologie
Universitätsstraße 150
D-44780 Bochum.
Terrestrial Isopoda from Guatemala and Mexico (Crustacea: Onisci-
dea: Crinocheta). - Some terrestrial Isopoda from Mexico, described in
the genus Philoscia Latreille, 1804, are redescribed and transferred to other
genera. Two new genera, Quintanoscia gen. n. and Oxalaniscus gen. n. are
described, they belong to the most primitive representatives of the Onis-
coidea within the Crinocheta. One of the species, Androdeloscia formosa
(Mulaik, 1960) is recorded from Guatemala for the first time. A closely
related species, A. valdezi sp. n. is described as new to science. New
records for Littorophiloscia denticulata (Ferrara & Taiti, 1981) and Burmo-
niscus kohleri (Schmalfuss & Ferrara, 1978) are presented; they were
recorded in the New World for the first time. Their distributional patterns
and phylogenetic relationships are discussed.
Key-words: Crustacea - Oniscidea - Philoscia - Androdeloscia - Littoro-
philoscia - Burmoniscus - new genera - new species - Central America -
taxonomy.
INTRODUCTION
Central America is one of the regions of the world, which is almost unexplored
with respect to the terrestrial isopod fauna. Only few contributions deal with taxa of
Oniscoidea collected in this biogeographically interesting area, as e.g., Miers (1877)
reporting on some Oniscidea from Central America. Particularly isopods of the
northern countries, Mexico and Guatemala, are little known. Richardson (1907)
described a new scleropactid species Spherarmadillo schwarzi Richardson, 1907,
from Guatemala and later a species of Armadillidae, Globarmadillo armatus Richard-
a Manuscript accepted 05.01.2000
284 ANDREAS LEISTIKOW
son, 1910 was described by Richardson (1910). This work remains the only contri-
bution to the Guatemalan oniscidean fauna.
The exhaustive contribution of Mulaik (1960) on the terrestrial isopod fauna of
Mexico is an important work, although the taxonomy used was no more up to date in
1960. Many species placed in the genus Philoscia Latreille, 1960 do not show any
apomorphic character in common with, e.g., the European Philoscia muscorum
(Scopoli, 1793), which is a typical member of this genus. Some of the species
included in Philoscia are among the most primitive representatives of Oniscoidea as
defined by Schmalfuss (1989): they, for instance, lack the noduli laterales. Some
species of Philoscia were revised in this study and transferred to other genera.
More recently, Schultz (1977) contributed to our knowledge of the troglobitic
species of Oniscidea from Central America, with Troglophiloscia laevis Schultz, 1977
being added to the Mexican fauna. The species is related to the Cuban 7. silvestrii
Brian, 1929, the type of the genus (Brian, 1929). The species might also occur in
northern Guatemala, as there are many caves in the northern province Petén, which
might be suitable habitats for this genus. However, it was not found in the Cueva
Actun Can near Santa Elena, Petén in a recently started survey.
This study aims to increase our knowlegde on the crinochete Oniscidea , which
are of philosciid appearance. Several species are described in detail to get access to a
complete data set for a phylogenetic analysis. New data are given for some species
probably introduced to Guatemala. Both Littorophiloscia denticulata (Ferrara & Taiti,
1981) and Burmoniscus kohleri (Schmalfuss & Ferrara, 1978) are recorded for the
first time from the New World. Ther distribtional patterns and phylogenetic relation-
ships are discussed.
The material examined is deposited in: Instituto Politécnico Nacional de
México (IPNM), Staatliches Museum für naturkunde (SMNS), Muséum d'histoire
naturelle de Geneve (MNHG), Universidad del Valle de Guatemala (UVG) and in the
author’s collection.
SPECIES ACCOUNT
Quintanoscia gen. n.
DIAGNOSIS: Cephalothorax without linea frontalis and linea supra-antennalis;
lamina frontalis present; compound eyes composed of 12 ommatidia; antennula
composed of three cylindrical articles; antennal flagellum three-articulate with apical
organ bearing two short free sensilla.
Mandibles with molar penicil consisting of three branches; medial endite of
maxillula with two penicils and apical tip: lateral endite with 4+6 teeth and slender
stalk; lobes of maxilla subrectangular: densely covered with trichiae; maxilliped
basipodite without sulcus lateralis; palp three-articulate; proximal article with two
long setae; medial and distal article with prominent setal tufts; endite elongate; setose;
with long penicil rostrally.
TERRESTRIAL ISOPODA FROM MEXICO AND GUATEMALA 285
Pereopods stout; antenna-grooming brush of carpus | arranged longitudinally;
dactylus with long inner claw; dactylar seta ending in a knob; coxal plates without
sulcus marginalis and gland pores; noduli laterales present; quite similar to tricorn-
like setae; insertion on all coxal plates at same distance from lateral margin.
Pleopods with rhomboidal exopodites bearing sensory spines laterally:
endopodites subquadrangular; no respiratory areas discernible at 400x magnification:
male genital papilla with ventral shield.
Uropod with protopodite laterally grooved; exopodite twice as long as the
more proximally inserted endopodite.
TYPE SPECIES: Philoscia contoyensis Mulaik, 1960 (by monotypy).
NUMBER OF NOMINAL SPECIES: only type species.
ETYMoLoGY: The genus is named after the Mexican province Quintana Roo, where
the type species was collected.
DISTRIBUTION: Only known from southeastern Mexico on the Yucatan Peninsula.
REMARKS: The genus Quintanoscia gen. n. is close to the groundpattern of the
Oniscoidea, the shape of the maxilliped is similar to Dero Guérin, 1836 and Allo-
niscus Dana, 1852. The pereopod 1 is equipped with a longitudinal carpal brush and
the medial margin is bearing several bifid sensory spines. The autapomorphies of the
genus are:
BM Reduction of the lateral lobes [lateral lobes present]
m Cephalothorax broadened [vertex not broader than height of cephalothorax]
The coxal plates bear several tricorn-like setae. One of those setae bears a
distinctly longer sensillum, which is twice as long as the basal cuticular plaque. This
structure can be interpreted as a nodulus lateralis.
Quintanoscia contoyensis (Mulaik, 1960) Figs 1-6
Philoscia contoyensis Mulaik, 1960
Material: Paratypes, 5 d (max. body length 3.5 mm): Mexico, Quintana Roo, Isla
Contoy, leg. 20.X1.1947, B.F. Osorio Tafall, IPNM 1631-A.
Colour: Mulaik (1960) wrote in the original description: “La coloracion de los
ejemplares conservados en alcohol; es un moteado de café; rojizo y armadillo. El
color mas obscuro esta confiando a una franja dorsal; media; delgada e irregular; a
una area angosta a lo largo de los margenes de os epimeros y entre estas; a otra banda
mas angosta fragmentada en manchas. La region ventral es casi blanca. Las formes
jovenes muestran menos pigmento que los adultos.”
Cephalothorax: Rather large with regard to body length; compound eyes
bearing 12 ommatidia; vertex strongly arched; no linea frontalis and linea supra-
antennalis visible; lamina frontalis inconspicuous (fig. 1; Ctf).
Pereon: Tegument smooth and shiny; bearing evenly spread tricorn-like setae;
noduli laterales similar to tricorn-like setae; sensillum comparatively longer; on all
coxal plates insertion at same distance to lateral margin; more distally located from
plate I to VII; no sulcus marginalis and gland pores.
286 ANDREAS LEISTIKOW
Pleon: Rather short; narrower than pereon; prominent neopleurae on pleonites
III to IV; pleotelson half as long than pleon; triangular; bordered by tricorn-like setae.
Appendages:
Antennula: Three-articulate with cylindrical articles; distal article bearing
longitudinally arranged aesthetascs (fig. 1; Anl).
Antenna: Peduncle covered with tricorn-like setae; flagellum three-articulate;
medial and distal article with pair of aesthetascs; apical organ as long as medial
article; with short free sensilla (fig. 5; An2).
Mouth parts as described in generic diagnosis (fig. 2).
Pereopods: Rather stout; carpus | with longitudinal antenna-grooming brush;
medial border of carpus and merus 1 to 4 bearing many bifid sensory spines (fig. 3-5;
PEI-7); dactylus with medium-sized inner claw (fig. 3; Dac); stout interungual seta;
tricorn-like seta laterally; dactylar seta with knob-like tip (fig. 4; Sd5). Sexual
dimorphism: Due to the lack of females not observed.
Pleopods: Exopodites rhomboidal with lateral margin bearing four to ten
sensory spines; on pleopod 4 and 5 one subapically on medial border; pleopod 5 with
small pectinate scales caudally; not arranged in rows; endopodites rounded triangular
to quadrangular; respiratory areas not discernible at 400x magnification (fig. 6; PLI-
5). Sexual dimorphism: Pleopod | endopodite long; rather prominent with ill-defined
basal area containing the intrinsic endopodite | levering muscle M49 (Erhard, 1997);
no row of small spines medio-caudally (fig. 6; PL1); pleopod 2 exopodite with lateral
margin slightly more sinuous than on exopodite 3; endopodite twice as long as
exopodite; rather stout (fig. 6; PL2).
Uropod: Protopodite triangular with lateral groove; endopodite inserting more
proximally than two times longer exopodite (fig. 5; UR).
Genital papilla: Ventral shield with parallel margins in basal half; slightly
surpassed by orifices (fig. 6; Gen).
Oxalaniscus gen. n.
DIAGNOSIS: Cephalothorax with lamina frontalis; linea supra-antennalis and
lateral lobes; linea frontalis reduced; compound eyes composed of about nine omma-
tidia. Antennula three-articulate; slender; antenna with three-articulate flagellum
(Mulaik, 1960).
Mandible with molar penicil composed of three branches; lateral endite of
maxillula with 4+5 simple teeth; slender stalk present; maxilla subrectangular;
maxilliped with long penicil on endite and prominent setal tufts on palp.
Pereopods with coxal plates lacking distinct nodulus lateralis; carpus 1 with
longitudinal carpal brush; setal brushes composed of sensory spines; not very dense;
present on carpus and merus | to 5; dactylar seta apically spatuliform.
Pleopods with prominent exopodites bearing few sensory spines laterally; no
respiratory structures discernible.
Uropod with protopodite subtriangular carrying lateral groove; endopodite
inserting slightly proximally of exopodite. Genital papilla with ventral shield sur-
passed by terminal spatula.
TERRESTRIAL ISOPODA FROM MEXICO AND GUATEMALA 287
/
EE —— 4
100um / li
Fic. 1: Quintanoscia contoyensis (Mulaik, 1960), 4 3.5mm body length. Anl antennula; Ctf
cephalothorax in frontal view; Cx4 coxal plate IV; Had habitus in dorsal view; Hal habitus in
lateral view; N14/7 setae of nodulus lateralis shape from coxal Figures IV and VII; Sx4 tricorn-
like seta of coxal plate IV; Tel pleotelson.
288 ANDREAS LEISTIKOW
SOum
Fic. 2: Quintanoscia contoyensis (Mulaik, 1960), 4 3.5mm body length. Mdl/r left and right
mandible; Mxp maxilliped, with detail of endite in rostral view; Mx1 maxillula, with details of
apical lateral endite in caudal and rostral view; Mx2 maxilla.
TERRESTRIAL ISOPODA FROM MEXICO AND GUATEMALA 289
100um
FIG. 3: Quintanoscia contoyensis (Mulaik, 1960), 3 3.5mm body length. Dac dactylus of pere-
opod I in rostral view; PE1-4 pereopods | to 4, in rostral (PE1) or caudal (PE2-4) view; Sbl
tricorn-like seta of basis 1; Scl sensory spine of carpal brush; Sc2 sensory spine of carpus 2.
290 ANDREAS LEISTIKOW
FIG. 4: Quintanoscia contoyensis (Mulaik, 1960), d 3.5mm body length. PES-6 pereopods 5
and 6 in caudal view; Sb6 tricorn-like seta of basis 6; Sc5/7 sensory spines of carpus 5 and 7;
Sd5 dactylar seta of dactylus 5.
TERRESTRIAL ISOPODA FROM MEXICO AND GUATEMALA 29]
FIG. 5: Quintanoscia contoyensis (Mulaik, 1960), d 3.5mm body length. An2 antenna, with
detail of apical organ; PE7 pereopod 7 in caudal view; UR uropod in rostral view.
ANDREAS LEISTIKOW
i)
Ne}
tO
FIG. 6: Quintanoscia contoyensis (Mulaik, 1960), 4 3.5mm body length. Gen genital papilla:
PL1-5 pleopods 1 to 5 in rostral view, with detail of endopodite 1 in caudal view.
TERRESTRIAL ISOPODA FROM MEXICO AND GUATEMALA 293
TYPE SPECIES: Philoscia ctenoscoides Mulaik, 1960 (by monotypy).
NUMBER OF NOMINAL SPECIES: Only type species included.
ETYMOLOGY: The genus name is derived from oxalä, the god of creation of the
indigenas from Brazil.
REMARKS: The genus Oxalaniscus gen. n. belongs to the most basal species of the
taxon Oniscoidea. It seems closely related to Quintanoscia gen. n. The putative
synapomorphies are given above. The following characters are the autapomorphies of
this genus:
BI Slender stalk of maxillula in a more lateral position [slender stalk medially,
surrounded by group of lateral teeth]
In all other taxa with a slender stalk on the maxillula, it is in a medial position
on the apical region, surrounded by the medial teeth of the outer group.
Oxalaniscus ctenoscoides (Mulaik, 1960) Figs 7-10
Philoscia ctenoscoides Mulaik, 1960
Material: several specimens: Mexico, Chiapas, Ruinas de Palenque, leg. 13.VII.1949,
C. and M. Goodnight, Finca Guautimoc, leg. 02.VII.1950, C. and M. Goodnight; Tabasco,
Emiliano Zapata, leg. 15.VIII.1945; Quintana Roo, Puerto Morelos, leg. XI.1947, B.F. Osorio
Tafall, Isla Cozumel, leg. 24.X1.1947, B.F. Osorio Tafall, IPNM 1628.
Colour: “Dorso brillante e intensamente moteado con café” (Mulaik, 1960).
Cephalothorax: Wider than high with linea supra-antennalis and lamina fron-
talis; small lateral lobes present; compound eyes composed of about nine ommatidia
(1 75 Cth):
Pereon: Tegument smooth with few tricorn-like setae; coxal plates without
gland pores; sulcus marginalis present; no noduli laterales visible (fig. 7, Cx4).
Pleon: Only slightly narrower than pereon; with prominent neopleurae on
pleonite 3 to 5; pleotelson with rounded apex; lateral margins straight.
Appendages:
Antennula: Slender; composed of three articles; distal one cylindrical with
apical pair of aesthetascs and medial set of about four aesthetascs (fig. 7, Anl).
Antenna: Peduncle with scattered tricorn-like setae; flagellum broken in all the
material examined; thus nothing can be said about the shape and the apical organ.
According to Mulaik (1960) it is composed of three articles.
Mouth parts as decribed in generic diagnosis (fig. 8).
Pereopods: Rather stout (fig. 9; PEI-7); carpus without latero-distal setal tuft;
no ornamental sensory spine on carpus |; antenna-grooming brush longitudinal;
carpus and merus | to 5 with loose brush of sensory spines; dactylus with short inner
claw (fig. 9, Dac); all interungual setae broken in material examined; dactylar seta
with flattened spatuliform apex (fig. 9, Sd1).
Pleopods: Exopodites of pleopod 3 to 5 large; bearing one to two sensory
spines laterally; no respiratory areas discernible in light microscope; endopodites
subrectangular (fig. 10, PLI-5). Sexual dimorphism: Male pleopod 1 exopodite
obtusely triangular; endopodite more than two times longer than exopodite; slightly
294 ANDREAS LEISTIKOW
bent sidewards; apex strongly bent sidewards with subterminal protrusions; about five
to ten small spines at proximal end of spermatic channel (fig. 10, PL1). Pleopod 2
exopodite triangular with single latero-distal sensory spine; distal third of endopodite
lace-shaped; slightly bent (fig. 10, PL2).
Uropod: As described in generic diagnosis.
Genital papilla: Ventral shield surpassed by terminally ending orifices; apex
indistinctly truncate (fig. 10, Gen).
REMARKS: The two new genera Quintanoscia and Oxalanoiscus from Mexico are
quite primitive with respect to several characters: They bear a subrectangular maxilla,
the maxillipedal palp is equipped with very prominent setal tufts and the penicil of the
endite is still prominent. Nonetheless, they are united in an sistergroup relationship
due to the following synapomorphies:
@ Cephalothorax with linea frontalis reduced [linea frontalis present]
@ Molar penicil composed of three branches [molar penixcil composed of
about ten branches]
@ Lateral endite of maxillula with 6+4 teeth, one of the inner set absent or at
least vestigial [no tooth reduced in size]
@ Maxillular teeth simple [teeth of inner set cleft]
In the outgroup, represented by Ligia baudiniana Milne Edwards, 1840, Deto
echinata Guérin, 1836 several species of Scleropatidae and Alloniscus Dana, 1852,
the characters differ considerably. The mandible is bearing a penicil of about ten
unfused branches and the maxillular teeth of the inner set are cleft or ctenate.
Therefore, the character states in both new genera are interpreted as being derived.
In spite of their close relationship, each species is placed in a monotypic genus
since the structure of the dorsal tricorn-like setae and the cephalothorax are striking.
Probably Quintanoscia and Oxalaniscus are as distant to each other as they are to
Alloniscus. In recent works on terrestrial isopods, the following characters are mostly
used for generic separation: cephalothorax morphology, structure and position of
dorsal receptors, presence of a sulcus marginalis, structure of the maxilliped,
arrangement of the teth on the lateral endite of the maxillula, setal patterns of the
pereopods, shape of pleon and pleotelson, shape of the male pleopod 5 and genital
papilla (Vandel, 1973a, 1973b; Taiti & Ferrara, 1980). Therefore, a generic separation
of the two taxa seems plausible.
Genus Littorophiloscia Hatch, 1947
DrAGNosis: Cephalothorax with linea supra-antennalis; slight lamina frontalis
and lateral lobes; linea frontalis lacking; compound eyes composed of about 15
ommatidia. Antennula three-articulate; slender; antenna long with three-articulate
flagellum; apical organ longer than distal article.
Mandible with molar penicil consisting of about ten free branches; maxillula
with lateral endite bearing 4+6 teeth; inner set with five teeth cleft; slender stalk pre-
sent; maxilla with both lobes subequal in width: maxilliped with endite setose; bea-
ring prominent penicil rostrally; palp three-articulate; with three prominent setal tufts.
TERRESTRIAL ISOPODA FROM MEXICO AND GUATEMALA 295
Cx4
Fic. 7: Oxalaniscus ctenoscoides (Mulaik, 1960), d 3mm body length. Anl antennula; Ctf
cephalothorax in frontal view; Cx4 coxal plate IV; Had habitus in dorsal view; Hal habitus in
lateral view; Ste tricorn-like seta of pleotelson; Tel pleotelson.
296 ANDREAS LEISTIKOW
Fic. 8: Oxalaniscus ctenoscoides (Mulaik, 1960), d 3mm body length. Mdl/r left and right
mandible; Mxp maxilliped, with detail of endite in rostral view; Mx1 maxillula, with detail of
apical lateral endite in rostral view; Mx2 maxilla.
TERRESTRIAL ISOPODA FROM MEXICO AND GUATEMALA 297
Na or LA
HERE Cares (EA)
GAL 7 \
Y &
STA
ge fy
| > 4
/
———
SOpim
Fic. 9: Oxalaniscus ctenoscoides (Mulaik, 1960), 4 3mm body length. Dac dactylus of
pereopod 1; PE1-7 pereopods | to 7 in rostral (PE1) or caudal view; Scl sensory spine of carpal
brush; Sd1 dactylar seta of pereopod 1.
298 ANDREAS LEISTIKOW
Fic. 10: Oxalaniscus ctenoscoides (Mulaik, 1960), ¢ 3mm body length. Gen genital papilla;
PL1-5 pleopods 1 to 5 in rostral view, with detail of endopodite 1 in caudal and rostral view.
TERRESTRIAL ISOPODA FROM MEXICO AND GUATEMALA 299
Pereopods slender; ornamental sensory spine of carpus 1 with serrate double-
fringe; dactylus with interungual seta long; more conspicuous than dactylar seta; inner
claw short; coxal plates without gland pores; sulcus marginalis and nodulus lateralis
present; all noduli inserted at same distance from lateral margin. Male pereopod 1
propus and carpus inflated; bearing more or less prominent setal brushes.
Pleopods without respiratory areas; exopodites with about five sensory spines
laterally; endopodites bilobate. Uropod with protopodite laterally grooved; endopodite
inserting more proximally than exopodite.
Genital papilla with short ventral shield; orifices on small to conspicuously
elongate protrusions of the terminal spatula.
TYPE SPECIES: Philoscia richardsonae Holmes & Gay, 1904.
NUMBER OF NOMINAL SPECIES: 23, four known from the Neotropics.
REMARKS: Taiti & Ferrara (1986a) revised the genus and gave a diagnosis for it, they
stressed the similarity of the male pereopods 1 and the mouth parts in Littorophiloscia
and Halophiloscia Verhoeff, 1908. With respect to the mouth parts, the polatity of the
characters was misinterpreted; they are of a plesiomorphic character state in both
genera, similar to the above described genera Quintanoscia and Oxalaniscus, and to
other Crinocheta with a position basal to the Oniscoidea sensu Schmalfuss (1989).
Littorophiloscia denticulata (Ferrara & Taiti, 1981) Figs 11-14
Bilawrencia denticulata Ferrara & Taiti, 1981
Material examined: 3 4,9 2 (ovigerous), 3 juveniles: Guatemala, Izabal, Livinston,
beach north of mouth of Rio Queguéche, between shells and fibres of coconuts, under Ipomoea
sp., leg. 22.X.1998, A. Leistikow, MNHG and author's collection.
Colour: Dorsally light violaceous brown with white paramedian patches; coxal
plates with two to three white patches; cephalothorax densely spotted with white;
pleon uniformly violaceous brown.
Cephalothorax: Linea frontalis reduced; linea supra-antennalis; lamina fron-
talis and small lateral lobes present; compound eyes comprising eight ommatidia (fig.
BIAC):
Pereon: Slightly arched; coxal plates with sulcus marginalis and nodulus
lateralis; distance between noduli and lateral margin subequal on all coxal plates (fig.
11, Cx4/Cxp); scattered tricorn-like setae present.
Pleon: Narrower than pereon; small neopleurae on pleonites 3 to 5 present;
pleotelson rounded with few tricorn-like setae along margin.
Appendages:
Antennula: Long and slender; three-articulate with pairs of aesthetascs in a
rowon distal article (fig. 11, Anl).
Antenna: Slender sparsely covered with of tricorn-like setae; flagellum three-
articulate with long apical organ; bearing short free sensilla; being as long as distal
article (fig. 11, An2).
Mandible: Pars intermedia with two penicils on left and one on right side;
bearing few setae; molar penicil composed of three branches; mandibular body
slender (fig. 12, Mdl/r).
300 ANDREAS LEISTIKOW
Maxillula: Medial endite with two weak penicils and apical tip; lateral endite
with 4+6 teeth; five of inner set cleft; slender stalk present (fig. 12, Mx1).
Maxilla: Subrectangular with both lobes subequal in width; scattered
trichiform setae present; more than ten cusps apically of medial lobe (fig. 12, Mx2).
Maxilliped: Basipodite slender; with sulcus lateralis; palp three-articulate; setal
tufts prominent; setae of proximal article subequal in length; endite slender; apically
setose; prominent penicil on rostral surface present (fig. 12, Mxp).
Pereopods: Slender; with scattered tricorn-like setae (fig. 13, PE1-7); carpus
with prominent antenna-grooming brush; along medial margin covering almost one
half of carpus length; ornamental sensory spine with hyaline apex; slightly striate (fig.
13, Scl); dactylus with simple dactylar seta (fig. 13, Dac); inner claw short; inter-
ungual seta with hyaline spatuliform apex. Sexual dimorphism: Male pereopods | to 2
with setal brushes on carpus; propus | inflated; equipped with prominent setal brush.
Pleopods: Pleopod exopodites rhomboid; with one to two sensory spines
laterally; endopodites subtriangular; no respiratory organs discernible in light micro-
scope (fig. 14, PLI-5). Sexual dimorphism: Male pleopod 1 endopodite rounded;
endopodite slender; more than two times longer than exopodite; apex acute; slightly
bent sidewards; distal third with caudomedial row of spines; laterally with three to
five subapical teeth (fig. 14, PL1). Pleopod 2 exopodite pointed; laterally sinuous;
endopodite slender; one third longer than exopodite (fig. 14, PL2).
Uropod: As in other species of the genus.
Genital papilla: Ventral shield with straight margins basally; terminal spatula
with orifices surpassing ventral shield; slightly bilobate (fig. 14, Gen).
DISTRIBUTION: Andaman Islands, Grub Island.
REMARKS: Littorophiloscia denticulata was described from the Andaman Islands as a
close relative of L. albicincta (Vandel, 1973) and L. occidentalis (Ferrara & Taiti,
1983). The outstanding autapomorphy of this species is the denticulated male pleopod
| endopodite; which is unique among its congeners. On the other hand, the species
mentioned above have a pleopod | with a similar structure. It is not possible to
evaluate the phylogenetic status of this character. Due to the relative simplicity of its
structure in comparison to its congeners, like L. richardsonae (Holmes and Gay,
1904) and L. riedlii (Strouhal, 1966), the latter formerly placed in Stenophiloscia
(Strouhal, 1966), L. denticulata may be close to the basis of this genus.
The occurence of this species in Guatemala most probably is due to human
introduction. The locality is 20 km north of Puerto Barrios, the main sea harbour of
Guatemala. L. denticulata may have been displaced with cargo from the South East
Asia, where it is assumed to be autochthonous, because its closest relatives occur in
the Indopacific region (Taiti & Ferrara, 1986a) and the radiation of this group might
have taken place in this area.
Burmoniscus Collinge, 1914
A diagnosis of Burmoniscus was given in Taiti & Ferrara (1986b).
TYPE SPECIES: Burmoniscus moulmeinus Collinge, 1914 (senior synonym of
Philoscia coeca Budde-Lund, 1895).
TERRESTRIAL ISOPODA FROM MEXICO AND GUATEMALA 301
250um
"n
100um
Fic. 11: Littorophiloscia denticulata (Ferrara & Taiti, 1981), 4 3.5mm body length. Anl
antennula; An 2 antenna, with detail of apical organ; Ctf cephalothorax in frontal view; Cx4
coxal plate IV, with detail of nodulus lateralis; Cxp coxal plates with position of noduli
laterales: Had habitus in dorsal view; Hal habitus in lateral view; Ste tricorn-like seta of
pleotelson; Tel pleotelson.
302 ANDREAS LEISTIKOW
|
Fic. 12: Littorophiloscia denticulata (Ferrara & Taiti, 1981), ¢ 3.5mm body length. Mdl/r left °
and right mandible; Mxp maxilliped, with detail of endite in rostral view; Mx! maxillula, with
details of apical lateral endite in rostral view; Mx2 maxilla.
TERRESTRIAL ISOPODA FROM MEXICO AND GUATEMALA 303
Fic. 13: Littorophiloscia denticulata (Ferrara & Taiti, 1981), d 3.5mm body length. Dac
dactylus of pereopod 1 in rostral view; PE1-7 pereopods 1 to 7 in caudal or rostral (PE1) view;
Sb4 tricorn-like seta of basis 4; Scl ornamental sensory spine of carpus 1.
304 ANDREAS LEISTIKOW
100um
RISI
___
I OOum — PL4
Fic. 14: Littorophiloscia denticulata (Ferrara & Taiti, 1981), 4 3.5mm body length /2 4.5mm
body length. Gen genital papilla; PEf female pereopod | in rostral view; PL1-5 pleopods 1 to 5
in rostral view, with detail of endopodite 1 in caudal view.
TERRESTRIAL ISOPODA FROM MEXICO AND GUATEMALA 305
NUMBER OF NOMINAL SPECIES: 58, two are reported from the Neotropics, both
of which most pobably introduced from West Africa and Southeast Asia.
DISTRIBUTION: Southeast Asia, East Africa, West Africa (?); some species with
circumtropical distribution due to dispersal by man.
REMARKS: The genus Burmoniscus Collinge, 1914 has been revised and newly
defined by Taiti & Ferrara (1986b). It is a heterogeneous, probably paraphyletic
group, comprising blind and unpigmented species and species which are superficially
similar to several members of /schioscia Verhoeff, 1928; e.g., Burmoniscus davisi
Taiti & Manicastri, 1988. In fact, no autapomorphies of the genus are described until
now. The characteristic position of the nodulus lateralis on coxal plate II 1s shared
with, e.g., Anchiphiloscia Stebbing; 1908 (Ferrara & Taiti, 1986), from which it
differs in the presence of gland pores and a sulcus marginalis, certainly plesio-
morphies of Burmoniscus.
The genus is distributed in the Oriental region, where it occurs with more than
30 species. The first record from South America was B. meussii (Holthuis, 1949),
reported from Brazil by Aratjo et al. (1996). It is found in suitable habitat in
Venezuela as well (pers. obs.). Records of the little known species Burmoniscus
kohleri (Schmalfuss & Ferrara, 1978) are presented from Guatemala for the first time.
Burmoniscus kohleri (Schmalfuss & Ferrara, 1978) Figs 15-18
Rennelloscia kohleri Schmalfuss & Ferrara, 1978
Material examined: 2 d (max.body length 2.5 mm), several 9: Guatemala, Izabal,
Aldea de Los Irrayoles, in scarse leaf litter in a plantation of Biepheridium? guatemalensis
(Rubiaceae), leg. A. Leistikow; 17.X.1998, MNHG and author's collection; 2 4 (max. body
length 2.5 mm); 5 9: Guatemala, Izabal, Aldea de Los Irrayoles; in a meadow with Cyperus
sp.; leg. 17.X.1998, A. Leistikow, author's collection; 2 4; 3 2: Guatemala, Izabal, Livinston,
finca in the western part of town on hill, under rotting stems of Cocos palms, leg. A. Leistikow;
21.X.1998; author's collection
Colour: Violaceous brown with light patches in medial line; paramedially and
on coxal plates; vertex densely spotted with light patches; pleon uniformly violaceous
brown.
Cephalothorax: Vertex slightly arched; compound eyes composed of about five
ommatidia; no linea frontalis and lamina frontalis; linea supra-antennalis present (fig.
1155 (Caop
Pereon: Slender; tegument smooth and shiny; scattered tricorn-like setae; coxal
plates with sulcus marginalis and nodulus lateralis; noduli of coxal plate II and IV
more remote from margin (fig. 15, Cx3/Cxp).
Pleon: Narrower than pereon; no neopleurae visible in dorsal view; pleotelson
with sinuous lateral margins; pointed; bearing few tricorn-like setae.
Appendages:
Antennula: Proximal article wide; medial and distal article forming a cone;
aesthetascs at top of distal article (fig. 15, Anl).
Antenna: Peduncle five-articulate; length ratio from proximal to distal joint 1:
2: 2: 3: 4; flagellum three-articulate with pairs of aesthetascs on medial and distal
article; apical organ as long as distal article; with short free sensilla (fig. 15, An2).
306 ANDREAS LEISTIKOW
SOum
Fic. 15: Burmoniscus kohleri (Schmalfuss & Ferrara, 1978), ¢ 2.7mm body length. Anl
antennula; An 2 antenna, with detail of apical organ; Ctf cephalothorax in frontal view; Cx3
coxal plate III; Cxp coxal plates with position of noduli laterales; Had habitus in dorsal view;
Hal habitus in lateral view; Tel pleotelson.
TERRESTRIAL ISOPODA FROM MEXICO AND GUATEMALA 307
Mandible: Molar penicil simple; pars intermedia with sparse setation; two
penicils on left; one on right side; intermedial penicil present (fig. 16, Mdl/r).
Maxillula: Medial endite pointed; bearing two penicils apically; lateral endite
with 4+5 teeth; three of inner set cleft; one double-cleft; one simple (fig. 16, Mx1).
Maxilla: Lateral lobe more than two times broader than medial one; setose;
laterally with transverse setal rows; medial lobe with few setae and five apical cusps
(fig. 16, Mx2).
Maxilliped: Basipodite with sulcus lateralis; palp with proximal article bearing
two setae; inserted close together; distal articles fused; two setal tufts present; pro-
ximal one with one prominent seta; endite with strong tooth caudally and knob-like
penicil rostrally (fig. 16, Mxp).
Pereopods: Slender with latero-distal setal tuft on carpus (fig. 17, PE1-7);
prominent transverse setal brush on rostral surface of carpus 1; half as broad as
medial margin; ornamental sensory spine double fringed (fig. 17, Sc1); dactylus with
short inner claw (fig. 17, Dac); dactylar seta simple; rather short. No sexual dimor-
phism discernible.
Pleopods: Pleopod exopodites rhomboid; bearing one to two sensory spines
laterally; single row of pectinate scales on caudal surface of exopodite 5; no
respiratory areas discernible; endopodites bilobate (fig. 18, PL1-5). Sexual dimor-
phism: Male pleopod 1 exopodite strongly concave on lateral margin; sinuosity
forming almost right angle; endopodite bent laterally; tapering in distal third; medio-
caudal row of short spines present (fig. 18, PL1). Pleopod 2 exopodite pointed;
laterally sinuous; endopodite slender (fig. 18, PL2).
Uropod: As in other species of the genus.
Genital papilla: Ventral shield stout; slightly surpassed by terminal spatula
with orifices (fig. 18, Gen).
DISTRIBUTION: West Africa, Cameroon.
REMARKS: This interesting species was found to be abundant in agricultural areas of
the Sierra de las Minas and the Caribbean coast at Livinston. They live in the leaf
litter or in rotten truncs of Cocos palms. B. kohleri was first discovered in Cameroon,
a remarkable location for a member of this genus; since all the other species described
are found in East Africa, South Asia and on the islands of the Wallacea. It is possible
that this species was also introduced to Cameroon, all the localities where the species
was found are close to the coast or to human settlements (Schmalfuss & Ferrara,
1978, 1983 and 1985). Thus, the natural occurence of this species may still be
unknown. On the other hand, Burmoniscus probably is paraphyletic and B. kohleri
might not belong to this genus. B. kohleri is quite distinct from its congeners due to its
minute size of maximally 3 mm and the number of spines along the medial margin of
carpus | is reduced to two as opposed to three to four spines in other species of
Burmoniscus.
If this species is native to Cameroon, it is quite isolated from the African
Oniscidean fauna by the position of the noduli laterales and the simple molar penicil,
with one exception: Taiti & Ferrara (1980 and 1982) listed for the West African area a
308 ANDREAS LEISTIKOW
Fic. 16: Burmoniscus kohleri (Schmalfuss & Ferrara, 1978), d 2.7mm body length. Mdl/r left
and right mandible, with detail of left pars intermedia; Mxp maxilliped, with detail of endite in
rostral view; Mx1 maxillula, with detail of apical lateral endite in rostral view; Mx2 maxilla.
TERRESTRIAL ISOPODA FROM MEXICO AND GUATEMALA 309
Fic. 17: Burmoniscus kohleri (Schmalfuss & Ferrara, 1978), & 2.7mm body length. Dac
dactylus of pereopod 7 in rostral view; PE1-7 pereopods | to 7 in caudal or rostral (PE1) view;
Sc] ornamental, longest and smallest sensory spines of carpus 1; Sp] medioproximal sensory
spine of propus 1.
310 ANDREAS LEISTIKOW
N
|
|
|
|
\
|
|
|
|
Fic. 18: Burmoniscus kohleri (Schmalfuss & Ferrara, 1978), d 2.7mm body length. Gen genital
papilla; PL1-5 pleopods 1 to 5 in rostral view, with detail of endopodite 1 in caudal view.
TERRESTRIAL ISOPODA FROM MEXICO AND GUATEMALA 311
genus which shares these characters. Zebrascia Verhoeff, 1942 was described on the
basis of a single female of 3 mm body length from Fernando Po (Verhoeff, 1942).
This genus now comprises three species; two of them from Cameroon (Schmalfuss &
Ferrara, 1978). Interestingly, the pereopod 1 carpus is of similar shape in both
Zebrascia and B. kohleri, and even the number and position of the sensory spines
correspond in both taxa. The position of the noduli laterales with respect to the
posterior margin of the coxal plates is similar in Zebrascia and B. kohleri (Schmalfuss
& Ferrara, 1978, Taiti & Ferrara, 1986b). It may be well possible that B. kohleri
belongs to a taxon close to Zebrascia; with the following synapomorphies:
M Antenna-grooming brush covering half the medial margin of pereopod
carpus | [antenna-grooming brush covering less than a third of carpus]
@ Sensory spines of carpus | arranged in a 2-spine-pattern [sensory spines
arranged in a 3-spine pattern]
In the Indopacific species of Burmoniscus differ from B. kohleri by the
pereopod | carpus being similar to the relations found in Androdeloscia (three-spine-
pattern, see below and fig. 20, PEI) and the antenna-grooming brush being not so
broad. To clarify this puzzling taxonomic situation, all species of Burmoniscus and
Zebrascia have to be re-examined.
Androdeloscia Leistikow, 1999
REMARK: The genus Androdeloscia was recently instituted for the small members of
the genus Prosekia Vandel, 1968; many species are found in northern South America
(Leistikow, 1999), but there are no records from Central America until now. How-
ever, two Central American species do belong to this genus, one originally described
as a member of Philoscia (Mulaik, 1960), the other a new species from Guatemala.
Beside /schioscia, Androdeloscia is the only genus of Crinocheta of philosciid appea-
rance known to occur in Central America and northern South America (Leistikow,
1997).
Androdeloscia formosa (Mulaik, 1960) Figs 19-22
Synonym: Philoscia formosa Mulaik, 1960
Material examined: 1 d (body length 3 mm); 1 9: Guatemala, Petén, El Remate,
Garden of the Casa de Don David, under coconuts and rotten leaves of Musa sp. and different
species of deciduous trees, close to pasture at Lake Petén Itza, leg. 27.X.1998, A. Leistikow,
author's collection; 3 4, several 2: Guatemala, Petén, causeway between Flores and Sta. Elena,
100 m off Ciudad de Flores; in rotten Eichhornia sp. directly at lake shore, leg. 30.X.1998, A.
Leistikow; author's collection and MNHG; 5 6, 4 ovigerous 9 : Guatemala, Petén, Sta. Elena,
road to Grutas Actin Can; 250 m from entrance, under bark of cut trees, leg. 30.X.1998; A.
Leistikow, author's collection; 3 d, 5 ® (Paratypes): Mexico, Tabasco, Palmillas, leg.
18.VIII.1945, F. Bonet, IPNM 1102-E.
Colour: Dorsally purplish brown with pale spots on pereon; medial line dark
brown; white central stripe on pereonites I-IV; double line on pereonites V-VII; pleon
unmarked.
312 ANDREAS LEISTIKOW
Cephalothorax: Linea frontalis and lamina frontalis lacking; linea supra-
antennalis present; lateral lobes small; compound eyes consisting of eight ommatidia
(fig. 19, Ctf).
Pereon: Tegument smooth and shiny; coxal plates without gland pores; sulcus
marginalis and noduli laterales present; the latter long and flagelliform; most dorsally
inserted on coxal plate IV (fig. 19, Cx3/Cxp).
Pleon: Narrower than pereon; neopleurae of pleonites 3 to 5 small; pleotelson
with straight distal margin; bearing some tricorn-like setae.
Appendages:
Antennula: Three-articulate with prominent proximal article; distal joint
bulbous; bearing two distinct sets of aesthetascs separated by a shallow depression
(fig. 19, Anl).
Antenna: Antennal peduncle composed of five articles with length ratio 1: 2: 2:
3: 4; densely covered with tricorn-like setae; flagellum composed of three articles;
distal one bearing prominent apical organ; as long as flagellar articles 1 and 2 together
(fig. 19, An2).
Mouth parts similar to following species.
Pereopods: Pereopods slender; with setal tuft latero-distally on carpus; carpus
1 with antennal-grooming brush; ornamental sensory spine serrate (fig. 20, Scl);
dactylus with short inner claw and simple dactylar seta; interungual seta straight (fig.
20, Dac). Sexual dimorphism: Male pereopod 7 merus with two small lobes medio-
distally; prominent lobe below distal sensory spine, directed distally; covered with
small scales (fig. 20, PE1-4; plate 21, PES-7).
Pleopods: Exopodites rhomboid with two sensory spines laterally; endopodites
subrectangular (fig. 22, PL1-5). Sexual dimorphism: Male pleopod 1 exopodite
rounded with slight concavity laterally; endopodite rather bulky; basal part containing
muscle M49 being half as broad as protopodite; laterally sinuous; distally with
prominent hump; apex slightly bent laterally; with short medio-caudal row of spines
groove for directing pleopod 2 exopodite; apex not elongate (fig. 22, PLS).
Uropod: As in other species of the genus.
Genital papilla: Rather short and slender compared to congeners (fig. 23, Gen).
REMARKS: This species belongs to a distinct subtaxon of the genus Androdeloscia
which is characterised by a sexual dimorphism of the male pereopod 7 merus. It bears
several hooks on its medial margin similar to A. silvatica (Lemos de Castro & Souza,
1986) and A. pseudosilvatica Leistikow, 1999 from the Caribbean region. Particularly
A. pseudosilvatica resembles A. formosa: the male pereopod 7 merus has three hooks
and the male pleopod | endopodite is bulbous at its base and falciform in the distal
part. Both species are separated by several characters. In A. formosa the maxillula has
a simple tooth more laterally, in A. pseudosilvatica the simple tooth is placed
medially, the male pereopod 7 merus bears two sensory spines in vicinity of the
proximal hook, in A. pseudosilvatica there is only one sensory spine, and the male
TERRESTRIAL ISOPODA FROM MEXICO AND GUATEMALA
313
200um
200m
Cxp
Fic. 19: Androdeloscia formosa (Mulaik, 1960), d 4mm body length. Anl antennula; An2
antenna, with detail of apical organ; Ctf cephalothorax in frontal view; Cxp position of noduli
laterales on coxal plates; Cx3 coxal plates3; Had habitus in dorsal view; Hal habitus in lateral
view; Tel pleotelson.
314 ANDREAS LEISTIKOW
Fic. 20: Androdeloscia formosa (Mulaik, 1960), ¢ 4mm body length. Dac dactylus 1 in rostral
view; PEI-4 pereopods 1-4 (caudal view), with detail of carpus 1 in rostral view; Scl orna-
mental sensory spine of carpus 1; Sc2 longest sensory spine of carpus 2; Spl distal sensory
spine of propus 1.
TERRESTRIAL ISOPODA FROM MEXICO AND GUATEMALA 315
Fic. 21: Androdeloscia formosa (Mulaik, 1960), ¢ 4mm body length. PES-7 pereopods 5-7 in
caudal view.
316 ANDREAS LEISTIKOW
PL2 PL3
Fic. 22: Androdeloscia formosa (Mulaik, 1960), 4 4mm body length. Gen genital papilla; PL1-
5 pleopods 1-5, rostral view, with detail of endopodite 1 in caudal view.
TERRESTRIAL ISOPODA FROM MEXICO AND GUATEMALA Se
pleopod 1 endopodite has the apex smooth and a lateral hump on half-length in A.
formosa, wheras in A. pseudosilvatica, the apex bears some lateral hooks and there is
no lateral hump. Additionally, the dactylar seta is acute in A. pseudosilvatica whereas
the apex of the dactylar seta in A. formosa is pointed. A. pseudosilvatica might be the
the adelphotaxon of A. formosa and A. silvatica, the latter share the lateral hump of
the male pleopod | endopodite as a synapomorphy.
Androdeloscia valdezi sp. n. Figs 23-26
Material examined: & (holotype, body length 3 mm): Guatemala, Petén, causeway
between Flores and Sta. Elena, 100 m off Ciudad de Flores, in rotten Eichhornia sp. at lake
shore, leg. 30.X.1998, A. Leistikow, MNHG uncatalogised; Paratypes: 3 4; several 2: same
data as for holotype, Guatemala, Petén, Sta. Elena, hill facing entrance to Grutas Actün Can,
scrubs, loamy soil sparsely covered with leaf litter, leg. 30.X.1998, A. Leistikow, author's
collection and UVG; 3 d, several 2: Guatemala; Zacapa, Rio Hondo, north of bridge over Rio
Hondo, right bank, in leaf litter, leg. 18.X.1998; A. Leistikow, SMNS.
DrAGNosis: Similar to most of its congeners, which do not show sexual dimorphism in
the pereopods; differs in the shape of male pleopod | endopodite, which is pointed
with lateral margin bearing several teeth.
Colour: Mostly as in other species of the genus; dorsally reddish brown with
light markings of muscle insertions; cephalothorax heavily spotted.
Cephalothorax: Linea frontalis missing; lamina frontalis weak; linea supra-
antennalis prominent; only slightly bent between antennal sockets; lateral lobes small;
compound eyes consisting of about eight ommatidia (fig. 23, Ctf).
Pereon: Tegument shiny; dorsum bearing scattered tricorn-like setae; coxal
plates with sulcus marginalis and flagelliform noduli laterales; the latter more dorsally
inserted on coxal plate IV (fig. 23, Cx3/Cxp).
Pleon: Retracted from pereon; neopleurae attached; pleotelson with straight
lateral margin; pointed; bearing short tricorn-like setae.
Appendages:
Antennula: Distal article apically with small tip and two aesthetascs; medially
with tuft of about nine aesthetascs; medial article distinctly shorter than proximal one
(fig. 23, Anl).
Antenna: More slender than in preceding species; especially flagellum; distal
article longest; proximal articles subequal in length; apical organ slender, longer than
distal article (fig. 23, An2).
Mandible: Molar penicil consisting of a three branches; pars intermedia
bearing two penicils on left and one on right mandible; intermedial penicil slender
(fig. 24, Mdl/r).
Maxillula: Medial endite similar to preceding species; lateral endite with 4+6
teeth; five of inner set cleft (fig. 24, Mx1).
Maxilla: Lateral lobe only slightly broader than medial lobe; bearing pectinate
scales; medial lobe sparsely covered with trichia; apically cuspidate (fig. 24, Mx2).
Maxilliped: As in other species of Androdeloscia; basipodite in examined
specimen broken (fig. 24, Mxp).
318 ANDREAS LEISTIKOW
Fic. 23: Androdeloscia valdezi sp. n., holotype & 3.5mm body length. Anl antennula; An2
antenna, with detail of apical organ; Ctf cephalothorax in frontal view; Cxp position of noduli
laterales on coxal plates; Cx3 coxal plate III; Had habitus in dorsal view; Hal habitus in lateral
view; Tel pleotelson.
TERRESTRIAL ISOPODA FROM MEXICO AND GUATEMALA 319
Fic. 24: Androdeloscia valdezi sp.n., holotype d 3.5mm body length. Mdl/r left and right
mandible, with detail of pars intermedia; Mxp maxilliped, with detail of endite in rostral view;
Mx1 maxillula, with detail of apex of lateral endite; Mx2 maxilla.
320 ANDREAS LEISTIKOW
Fic. 25: Androdeloscia valdezi sp. n., holotype & 3.5mm. Dac dactylus 1 in rostral view:
PEI-7 pereopods 1 to 7 (caudal view), with details of carpus 1 in rostral and ischium 6 in
caudal view; Scl ornamental sensory spine of carpus 1: Sp! distal sensory spine of propus 1.
TERRESTRIAL ISOPODA FROM MEXICO AND GUATEMALA 32]
FIG. 26: Androdeloscia valdezi sp. n., holotype 4 3.5mm body length. Gen genital papilla;
PLI-5 pleopods 1-5, rostral view, with details of endopodite 1 in caudal and rostral view.
322 ANDREAS LEISTIKOW
Pereopods: Rather slender; especially pereopod 1; carpus with antenna-groo-
ming brush and serrate ornamental sensory spine (fig. 25, Sc1); pereopod 2 merus
with hyaline fringe medially; dactylus with long inner claw and simple dactylar seta
(fig. 25, Dac). Sexual dimorphism not evident (fig. 25, PE1-7).
Pleopods: Pleopod exopodites 3 and 4 elongate rhomboid with two sensory
spines laterally; exopodite 5 more or less triangular; endopodites bilobate; no respira-
tory areas discernible in light microscope (fig. 26, PL1-5). Sexual dimorphism: Male
pleopod 1 exopodite rounded; endopodite pointed with lateral margin bearing several
teeth; basal part with muscle M49 less than half the length of endopodite; short caudal
row of sensory spines (fig. 26, PL1). Pleopod 2 and 5 similar to preceding species
(He227APE2/5);
Uropod: As in other species of the genus.
Genital papilla: Similar to preceding species (fig. 26, Gen).
REMARKS: The new species is close to the “species-group D” of the genus Andro-
deloscia with the autapomorphies: hyaline lobes on the male pleopod 1 endopodite
reduced; maxillipedal basipodite slender, apically subrectangular (Leistikow, 1999).
The exact systematic position within this group is difficult to access: the male pleopod
1 endopodite in A. valdezi is similar to that of A. conipus Leistikow, 1999 and related
species, but this similarity is possibly a symplesiomorphy. A. poeppigi Leistikow,
1999 and A. malleus Leistikow, 1999 form the sister group of A. conipus and related
species. The former have the more derived endopodites | within species-group D. The
caudo-medial row of spines is reduced and there are two subapical lobes, synapo-
morphic characters of A. poeppigi and A. malleus. For the time being, the systematic
positon of the new species remains open to debate.
ACKNOWLEDGEMENTS
The author wishes to thank the following persons: Prof. Dr J.W. Wägele,
Ruhr-Universität Bochum, for critically discussing the results of this work; Dr M.
Dix, Mrs Mayra Maldonado and Mr José Rodolfo Valdez Barilla, Universidad del
Valle de Guatemala and Mrs. Beate Fasselt for their support during the field trip in
Guatemala. For financial support of the field trip, the author is indebted to the
Studienstiftung des Deutschen Volkes. This study is dedicated to the victims of the
Hurricane “Mitch”, which devastated the northern Central American region in
October 1998.
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REVUE SUISSE DE ZOOLOGIE 107 (2): 325-334; juin 2000
Euphalerus clitoriae sp. n., a new psyllid species from
Clitoria fairchildiana (Fabaceae, Papilionoideae), and notes
on other Euphalerus spp. (Hemiptera, Psylloidea)
Daniel BURCKHARDT! & Mariängela GUAJARA?
| Naturhistorisches Museum, Augustinergasse 2, CH-4001 Basel, Switzerland.
E-mail: daniel.burckhardt@unibas.ch
2 Universidade Federal Rural do Rio de Janeiro, CPG em Fitotecnia
Seropédica, RJ — 23.851-970, Brazil.
E-mail: guajara@ufrrj.br
Euphalerus clitoriae sp. n., a new psyllid species from Clitoria fair-
childiana (Fabaceae, Papilionoideae), and notes on other Euphalerus
spp. (Hemiptera, Psylloidea). - Euphalerus clitoriae sp. n. from Brazil, a
species developing under waxy filamentous secretions on the leaves of
Clitoria fairchildiana, ıs described and illustrated. Its biology is briefly
outlined and compared to that of its closest relatives, viz. E. nidicola from
Peru and E. maya from Belize. Differences to these two species are
discussed. Based on the study of types, the genitalia of E. nidicola, E.
antillensis and E. ostreoides are illustrated completing their insufficient
original descriptions.
Key-words: Psylloidea - Fabaceae - Brazil - taxonomy - biology.
INTRODUCTION
The hemipterous jumping plant-lice or psylloids are usually highly host specific
and thus a potentially suitable taxon for coevolutionary studies on insect — plant
relationships (Burckhardt & Basset, 2000). The necessary taxonomic and phylogenetic
base is, however, often insufficient or completely lacking. The genus Euphalerus is a
good example. It was erected for the Carribean species E. nidifex whose larvae form
lerps or “nests”, i.e. waxy coverings, on Fish poison Bark or Jamaican dogwood
(Piscidia piscipula and Piscidia carthegenensis, Fabaceae, Papilionoideae, Tephro-
sieae) (Schwarz, 1904; Russell, 1971). Subsequently, a series of mostly tropical New
and Old World species were referred to the genus making it extremely artificial.
According to Hollis & Martin (1997), the New World species forming lerps or galls on
the leaves of Fabaceae make up true Euphalerus (Psyllidae, Euphalerinae). The other
New World species, associated with Ceanothus (Rhamnaceae) and Cerocarpus (Rosa-
ceae), are unrelated and belong to the Arytaininae/Psyllinae (Psyllidae) complex. The
Old World species, finally, are probably referable to the euphalerine genera Colo-
phorina Capener and Euryconus Aulmann.
Manuscript accepted 08.11.1999
326 DANIEL BURCKHARDT & MARIANGELA GUAJARA
Twenty two species are currently known in Euphalerus s. str. The vast majority
is associated with Lonchocarpus (Papilionoideae, Tephrosieae). One species is each on
Pithecellobium (Mimosoideae, Ingeae), Erythrina (Papilionoideae, Phaseoleae) and
Piscidia (Papilionoideae, Tephrosieae) respectively, two species have unknown and one
doubtful (Karwinskia, Rhamnaceae) host records. Nineteen species are recorded from
Central America with one species extending to Florida, and only three are also or
exclusively known from South America (Brazil 2, Peru 1) (Hollis & Martin, 1997).
The present paper describes a new species from Brazil developing on Clitoria
fairchildiana (Papilionoideae, Phaseoleae). The new species broadens the known host
range of Euphalerus, and suggests that the genus may be more diverse in tropical South
America than currently estimated. It is closely related to the gall-forming E. nidicola
Tuthill from Peru whose larvae are covered in waxy secretions, and the lerp-inhabiting
E. maya Hollis & Martin from Belize. Hollis & Martin (1997) provided detailed
descriptions of the species attacking Lochocarpus in Belize. Here we give illustrations
of the male and female genitalia of E. antillensis Caldwell & Martorell and E. nidicola
Tuthill, and of the male genitalia of E. ostreoides Crawford whose original descriptions
lack sufficient detail and which were not treated by Hollis & Martin (1997).
MATERIAL AND METHODS
The morphological terminology follows Hollis (1976) and Hollis & Martin
(1997). Measurements and some drawings were made from slide mounted material. The
drawings for E. antillensis and E. nidicola were made from temporary mounts in
glycerine.
Material is cited from following institutions: Naturhistorisches Museum Basel,
Switzerland - NHMB; Angelo Moreira da Costa Lima Entomological Collection, Sero-
pédica, Brazil - CLEC; Natural History Museum, London, UK - BMNH; United States
National Museum of Natural History, Washington, DC, USA (psylloid collection in
USDA, Beltsville, MD) - USNM; Muséum d’histoire naturelle, Geneva, Switzerland -
MHNG.
TAXONOMIC TREATMENT
Euphalerus clitoriae sp. n. Figs 1-3, 13, 16-23
Holotype & , Brazil: State of Rio de Janeiro, Campus and residential area of the Univer-
sidade Federal Rural do Rio de Janeiro, Seropédica, 22°44' S 43°43' W, 9.1x.1999, Clitoria
fairchildiana (M. Guajara), dry mounted (NHMB).
Paratypes, Brazil: 20 d, 20 9, 10 larvae or larval skins, same data as holotype, dry
mounted (NHMB, CLEC, BMNH, USNM, MHNG); 6 d, 6 ©, 4 larvae, same data but 30.v.
1999, dry and slide mounted (NHMB, CLEC, BMNH).
Material not included in type series. Brazil, many adults and some larvae, same data as
holotype but 30.v.1999 and 9.1x.1999, preserved in 70 % alcohol (NHMB).
Adult. Body dirty greyish white with small blackish dots, in mature specimens
thorax dorsally, abdomen and genitalia dark. Antennal segments 1-8 light with dark
apices, segments 9 and 10 entirely dark. Forewing membrane transparent, colourless,
with blackish dots as in fig. 16, middle of cells very weakly infuscate; veins whitish
with black dots. Legs whitish with dark dots, tarsi blackish.
EUPHALERUS CLITORIAE 327
Small species (see measurements below). Genal processes longer than vertex
along mid-line, conical, subacute apically (fig. 17). Antenna 1.97-2.13 times as long as
head width, segment 3 (1.32 times) longer than segment 8. Ultimate two rostral
segments 0.41-0.43 times as long as head width. Forewing (fig. 16) 2.24-2.31 times as
long as wide, 3.67-3.93 times as long as head width; pterostigma moderately long, 0.5
times as long as vein Rs; surface spinules leaving broad spinule-free stripes along the
veins; reduced in basal half of cell r, and r,, occupying only a small area in cell c+sc.
Metatibia 0.93-0.97 times as long as head width, bearing a small basal spine and an
incomplete crown of apical spurs.
Male proctiger (fig. 1) bulbous, lacking lateral lobes. Paramere in profile (fig. 2)
with each an anterior and a posterior lobe. Distal portion of aedeagus (fig. 3) hook-
shaped; apex of terminal tube of ductus ejaculatorius truncate.
Female proctiger (fig. 13) more or less evenly tapering, dorsal margin weakly
concave, apex narrowly rounded; 1.00 times as long as head width, 3.21 times as long
as circumanal ring, 1.56 times as long as subgenital plate.
Measurements in mm (1 6, 1 9). Head width 0.63; antenna length 1.25-1.35;
forewing length 2.33-2.50; male proctiger length 0.26; paramere length 0.23; length of
distal segment of aedeagus 0.23; female proctiger length 0.63.
Fifth instar larva (fig. 18). Antenna 8-segmented with one rhinarium on each of
segments 3, 5, 7 and 8. Tibiatarsus of foreleg with one large apical and one smaller
subapical spur (fig. 19); tibiotarsi of mid and hindlegs with one large and two small
spurs. Subgenital plate with two apical processes bearing three teeth each (fig. 20).
Host plant. Clitoria fairchildiana Howard (= racemosa) (Fabaceae).
Biology. The eggs are laid along the veins of the leaves. The larvae sit on the
leaves and stems hidden under white coverings consisting of waxy hair-like filaments
(fig. 21). There is no sign of deformations on the leaves (fig. 22). When the larval
density is high, the leaves turn yellow and drop. Strongly infested trees can loose all
their leaves (fig. 23).
Comments. E. clitoriae, which is a member of the E. ostreoides species group as
defined by Hollis & Martin (1997), is closely related to E. nidicola Tuthill, 1959 from
Peru, and E. maya Hollis & Martin, 1987 from Belize based on following characters.
Small sized Euphalerus spp.; head with long, conical genal processes; antenna longer
than 1.5 times head width; forewing clear, bearing dark spots lacking transverse bands;
metatibia with a small basal spine and an incomplete crown of apical spurs; male
proctiger bulbous without lateral lobes; paramere complex, irregularly S-shaped with a
group of long thick setae in the middle of the foremargin and a group of heavily
sclerotised peg setae on the inner surface of the posterior lobe; female subgenital plate,
in profile, with angular ventral margin.
E. clitoriae differs from E. maya in the slightly larger body dimensions, the
longer antenna, the shorter pterostigma of the forewing, the details in the genital
structure and the apex of the larval caudal plate which has three points on each tubercle
instead of two. E. clitoriae is differentiated from E. nidicola by the much more
expanded, almost black body coloration, which is orange to light brownish in E.
nidicola, the blackish dots on the forewings which are brown in E. nidicola, the shorter
328 DANIEL BURCKHARDT & MARIANGELA GUAJARA
Fics 1-6
Euphalerus spp. 1, 4: Male genitalia, in profile; 2, 5: paramere, inner face; 3, 6: distal portion of
aedeagus. 1-3: E. clitoriae sp. n.; 4-6: E. nidicola Tuthill. Scale lines: figs 1, 4 = 0.2 mm, figs 2,
3, 4,6=0.1 mm.
EUPHALERUS CLITORIAE 329
0.2 mm
0.3 mm
Fics 7-12
Euphalerus spp. 7, 10: Male genitalia, in profile; 8, 11: paramere, inner face; 9, 12: distal portion
of aedeagus. 7-9: E. antillensis Caldwell & Martorell; 10-12: E. ostreoides Crawford. Scale lines:
fig. 7 = 0.2 mm, figs 8, 9 = 0.1 mm, fig. 10 = 0.3 mm, figs 11, 12 = 0.2 mm.
antenna, the more reduced fields of surface spinules in the forewing, the details of the
genitalia (for E. nidicola see figs 4-6, 14); the female proctiger is slightly shorter and
apically more thickened in E. clitoriae but longer and apically pointed in E. nidicola.
The three species differ also in their geographical and host plant ranges. The
larval biology of E. clitoriae is intermediate. It shares with E. nidicola waxy filamen-
330 DANIEL BURCKHARDT & MARIANGELA GUAJARA
tous coverings but differs by not causing depressions on the leaves. This character is as
in E. maya which is characterised by the production of a lerp.
Euphalerus antillensis Caldwell & Martorell Figs 7-9, 15
Euphalerus antillensis Caldwell & Martorell, 1951: 612. Holotype d, Puerto Rico: Guänica, on
the Guanica - Guänica Central Road, in front of stone quarry, 28.viii.1947, breeding abundantly
on the undersides of the leaves of “geno-geno”, Lonchocarpus domingensis (Caldwell &
Martorell) (USNM).
Material examined. Puerto Rico: 1 d, 1 2 paratypes, same data as holotype (USNM).
Adult. Member of the nidifex species group (as defined by Hollis & Martin,
1997). Metatibia with a small basal spine and grouped apical spurs.
Male proctiger (fig. 7) with narrow lateral lobes. Paramere in profile (fig. 8)
lamellar, obliquely truncate apically, inner surface covered in long setae. Distal portion
of aedeagus (fig. 9) with oval apical dilatation, rounded apically; apex of terminal tube
of ductus ejaculatorius rounded.
Female proctiger (fig. 15) strongly tapering to the middle, digitiform in apical
half, dorsal margin strongly concave, apex narrowly rounded.
Euphalerus nidicola Tuthill Figs 4-6, 14
Euphalerus nidicola Tuthill, 1959: 6. Holotype d, Peru: few km below La Merced, 1.1.1947, on
“oropel”, Erythrina sp. (Tuthill) (USNM).
Material examined. Peru: holotype à , allotype 2,1 d,1 © paratypes (USNM).
Adult. Member of the ostreoides species group (as defined by Hollis & Martin,
1997).
Male proctiger (fig. 4) bulbous, lacking lateral lobes. Paramere in profile (fig. 5)
with each an anterior and a posterior lobe. Distal portion of aedeagus (fig. 6) hook-
shaped; apex of terminal tube of ductus ejaculatorius truncate.
Female proctiger (fig. 14) irregularly tapering, dorsal margin weakly but
distinctly concave, apex subacute.
Fifth instar larva. Tuthill (1959) mentioned the presence of many larvae. In the
collection of the USNM we could, however, find only the adult types.
Euphalerus ostreoides Crawford Figs 10-12
Psyllid, Tavares, 1920: 124; 1922: plate 19 figs 3-5. Galls on the leaves of an undetermined
species of Leguminosae (Timb6), Brazil: Rio de Janeiro State, Nova Friburgo; Rio de Janeiro
State, surroundings of Rio de Janeiro; Sao Paulo State, Itu; between Rio Vermelho and the city of
Bahia.
Euphalerus ostreoides Crawford, 1925: 62. Syntypes 2 d, 1 ®, Brazil: Sao Paulo, Itu, making
very peculiar galls on the leaves of an undetermined species of Leguminosae (Tavares) (USNM).
Costa Lima, 1942: 107; Silva et al., 1968: 201; Russell, 1971: 10; Hollis & Martin, 1997: 241.
Material examined. Brazil: syntype d, Sao Paulo, Itu, (Tavares) (slide mounted, USNM).
Adult. Member of the ostreoides species group (as defined by Hollis & Martin,
1997). Large species (see measurements below). Genal processes shorter than vertex
along mid-line, broadly rounded apically. Antenna 2.32 times as long as head width,
segment 3 about as long as segment 8. Ultimate two rostral segments 0.39 times as long
EUPHALERUS CLITORIAE 331
=
Ve 4,
0.2 mm a >
“2 7; "pt
Ni i fe
Fics 13-15
Euphalerus spp., female genitalia, in profile. 13: E. clitoriae sp. n.; 14: E. nidicola Tuthill; 15:
E. antillensis Caldwell & Martorell. Scale line = 0.2 mm.
332 DANIEL BURCKHARDT & MARIANGELA GUAJARA
c
E
e
ei
a
Fics 16-20
Euphalerus clitoriae sp. n. 16: forewing; 17: head, dorsal view; 18: fifth instar larva, left dorsal,
right ventral view; 19: apex of foreleg of fifth instar larva; 20: apex of caudal plate, dorsal view.
Scale lines: fig. 16 = 1.0 mm, fig. 17 = 0.3 mm, fig. 18 = 0.3 mm, figs 19, 20 = 0.1 mm.
333
EUPHALERUS CLITORIAE
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334 DANIEL BURCKHARDT & MARIANGELA GUAJARA
as head width. Forewing 2.34 times as long as wide, 3.06 times as long as head width;
pterostigma relatively short, 2.6 times as long as vein Rs; surface spinules present in
apical part of all cells except for c+sc, leaving broad spinule-free stripes along the
veins; radular spinules covering broad triangular patches along the wing margin in cells
Ty, 1, Mj, m, and cu,. Metatibia 0.77 times as long as head width, bearing a conspi-
cuous basal spine and an incomplete crown of apical spurs.
Male proctiger (fig. 10) with distinct lateral lobes. Paramere in profile (fig. 11)
with a large posterior lobe, inner face with an anterior field of longer, more spaced setae
and a posterior field with peg-like, densely spaced setae. Distal portion of aedeagus
(fig. 12) rounded apically, with inflated apical half; apex of terminal tube of ductus
ejaculatorius truncate.
Female unavailable.
Measurements in mm (1 9). Head width 1.15; antenna length 2.68; forewing
length 3.54; male proctiger length 0.47; paramere length 0.47; length of distal segment
of aedeagus 0.47.
ACKNOWLEDGEMENTS
We thank D. Miller (USDA, Beltsville Md, USA) for the loan of material, D.
Hollis (BMNH) for comments on Euphalerus and on a manuscript draft, and A. G. de
Carvalho (Seropédica) for pointing out the new species and suggesting its study.
REFERENCES
BURCKHARDT, D. & BASSET, Y. 2000. The jumping plant-lice (Hemiptera, Psylloidea) associated
with Schinus (Anacardiaceae): systematics, biogeography and host plant relationships.
Journal of Natural History 34: 57-155.
CALDWELL, J. S. & MARTORELL, L. F. 1951. A brief review of the Psylliidae of Puerto Rico
(Homoptera). Annals of the Entomological Society of America 44: 603-613.
Costa LIMA, A. DA. 1942. Homopteros. Superfamilia Psylloidea. Insetos do Brasil 3: 94-111.
CRAWFORD, D. L. 1925. Psyllidae of South America. Broteria 12: 56-74.
HoLLis, D. 1976. Jumping plant lice of the tribe Ciriacremini (Homoptera: Psylloidea) in the
Ethiopian Region. Bulletin of the British Museum (Natural History) (Entomology) 43:
1-83.
Ho tis, D. & MARTIN, J. 1997. Jumping plantlice (Insecta: Hemiptera) attacking Lonchocarpus
species (Leguminosae), including ‘Black Cabbage Bark’, in Belize. Journal of Natural
History 31: 237-267.
RUSSELL, L. M. 1971. Notes on Euphalerus nidifex Schwarz and related nest-making New World
psyllids (Homoptera: Psyllidae). Florida Entomologist 54: 3-12.
SCHWARZ, E. A. 1904. Notes on North American Psyllidae. Part I. 2. Description of the nest-
constructing Psyllid. Proceedings of the entomological Society of Washington 6: 234-245.
SILVA, A. G. D’ ARAUJO E, GONÇALES C. R., GALVAO D. M., GONGALES A. J. L., GOMES J., SILVA,
M. DO N. & SIMONI, L. DE. 1968. Quarto catalogo dos insetos que vivem nas plantas do
Brasil, seus parasitos e predadores. Parte II - I tomo, insetos, hospedeiros e inimigos
naturais, Rio de Janeiro, xxiv + (iv) + 622 pp.
TAVARES, J. S. 1920. Cecidologia Brazileira. Broteria, Série Zoologica 18: 82-125.
TAVARES, J. S. 1922. Cecidologia Brazileira. Broteria, Série Zoologica 20: 5-48.
TUTHILL, L. D. 1959. Los Psyllidae del Peru Central (Insecta: Homoptera). Revista Peruana de
Entomologia Agricola 2: 1-27.
REVUE SUISSE DE ZOOLOGIE 107 (2): 335-349; juin 2000
Zur Biologie des “Eingeweidefisches”
Carapus acus (Brünnich, 1768) (Carapidae, Teleostei),
mit Hinweisen auf eine nicht-parasitische Ernährung!
Kerstin KLOSS und Wolfgang PFEIFFER
Zoologisches Institut der Universität Tübingen
Auf der Morgenstelle 28, D-72076 Tübingen,
und
Laboratoire Arago, Universite Paris VI
F - 66650 Banyuls-sur-mer.
On the biology of the pearlfish Carapus acus (Brünnich, 1768) (Cara-
pidae, Teleostei), with indications of a non-parasitic nutrition. - 450
Stichopus regalis (Cuvier), caught with a trawl-net at a depth of 70-110 m
off Banyuls-sur-mer (France) contained 38 Carapus acus. We found only
two pearlfish in 85 Holothuria tubulosa Gmelin caught in less than 10 m
of seawater. We kept pearlfish in aquaria and analysed their behaviour by
means of computer analysis of video records. The manner by which the
fish enter their hosts was studied (see summary). In the wild most spe-
cimens of C. acus are found in S. regalis, yet in our aquaria C. acus only
entered H. tubulosa if they could choose between both species. We studied
the stomach contents of C. acus and found crustaceans. The stomachs
examined did not contain any holothurian tissue. Our results give reason to
agree with Emery (1880), who stated that C. acus is an inquiline, to whom
the host holothurian serves as a shelter only.
Key-words: pearlfish - Carapus acus - Carapidae - Teleostei - host holo-
thurian - Stichopus regalis - Holothuria tubulosa - stomach contents -
behaviour - parasitism - inquilinism.
EINLEITUNG
Der “Eingeweidefisch” Carapus acus (Briinnich, 1768) (gängiges Synonym
Fierasfer acus) ist in jeder Beziehung außergewöhnlich und erregte aufgrund seiner
besonderen epidermalen Kolbenzellen bereits 1957 die Aufmerksamkeit des Zweit-
Autors (W. P.). Diese Zellen ähneln weitgehend den Schreckstoffzellen der Ostario-
physi (Pfeiffer, 1960). Karl von Frisch, Doktorvater des Zweit-Autors, forderte diesen
bei seiner Abreise an die Zoologische Station in Neapel Anfang 1957 auf, den
Mageninhalt von C. acus zu untersuchen. Dies geschah an 2 von Pierre Tardent zur
! In memoriam Prof. Dr. Pierre Tardent.
Manuskript angenommen 09.11.1999
336 KERSTIN KLOSS UND WOLFGANG PFEIFFER
Verfügung gestellten Exemplaren; beide enthielten Reste von Kleinkrebsen, wie an den
schwarzen Komplexaugen deutlich zu erkennen war. Dieser Befund wurde Alfred
Kaestner mitgeteilt und ist in dessen Lehrbuch nachzulesen (Kaestner, 1963, S. 1235).
Zwanzig Exkursionen an das Laboratoire Arago in Banyuls-sur-mer (1963 und 1980 -
1999) ermöglichten C. acus zu beobachten, zu sammeln und mit ihm zu experimen-
tieren. Im Mittelpunkt des Interesses standen dabei die Fragen nach seiner Häufigkeit
und Verbreitung, seinem Habitat und der Bevorzugung verschiedener Seewalzen-
Species, der Art und Weise seines Eindringens, der Verbleibdauer im Wirt und nicht
zuletzt seiner Ernährung. Handelt es sich bei C. acus tatsächlich um einen Nahrungs-
parasiten, der Teile der Seewalze frißt, wie von Arnold (1953) behauptet aber nicht
bewiesen und von der Sekundarliteratur kritiklos übernommen worden ist? Oder ist C.
acus ein Inquilinist, dem die Seewalze lediglich als Wohnhöhle dient, wie von Emery
(1880) beschrieben? Diesen Fragen sind wir im folgenden nachgegangen.
MATERIAL UND METHODEN
Alle Exemplare von C. acus wurden an der Küste vor Banyuls-sur-mer (Frank-
reich) gesammelt: 38 C. acus stammten aus 450 Stichopus regalis (Cuvier) (70 - 110 m
Meerestiefe), nur 2 aus 85 Holothuria tubulosa Gmelin (weniger als 10 m Meerestiefe).
Demgegenüber enthielten 142 Holothuria polii Delle Chiaje und 63 H. tubulosa aus der
Bucht von Port de la Selva (weniger als 10 m Wassertiefe) keinen einzigen C. acus.
Weil die Fische bisher fast ausschließlich in ihren Wirten und nahezu nie frei-
schwimmend gefunden worden waren, konzentrierten wir uns beim Fang auf die als
Wirte bekannten Aspidochirota S. regalis, H. tubulosa und H. polii. Die Seewalzen
wurden mittels eines Schleppnetzes gesammelt, das in 70 - 110 m Tiefe 60 - 90 min
lang parallel zur Tiefenlinie über den Grund gezogen wurde. Die dabei gefangenen
S. regalis - die beiden anderen Species kommen in dieser Tiefe vor Banyuls-sur-mer
nur äußerst selten vor - wurden an Bord in einem Behälter mit Meerwasserdurchlauf
aufbewahrt und an Land aufgeschnitten, um zu prüfen ob sie C. acus enthielten. Beim
Eröffnen ihrer physiologischen Unterseite mit einer kräftigen Präparierschere war es
unerheblich, ob der Schnittansatz im Maul oder im After erfolgte. Die Fische können
sich aufgrund ihrer flexiblen Wirbelsäule an dem der Einschnittstelle gegenüber-
liegenden Ende zu einem kleinen Knäuel zusammenrollen, so daß sie leicht zu
übersehen sind (Abb. 2e). Wegen des grobmaschigen Schleppnetzes war nicht gewähr-
leistet, daß freischwimmende C. acus oder solche, die von ihrer Seewalze mitsamt den
Eingeweiden ausgestoßen worden waren, mitgefangen wurden. Um das Entkommen
der Fische zu vermeiden, wurden H. tubulosa und H. polii von Hand gesammelt und
unter Wasser in Plastiksäcken aufbewahrt. Beim Aufschneiden war erhöhte Vorsicht
geboten, da die Arten von Holothuria eine wesentlich stärkere Hautmuskulatur besitzen
als S. regalis und sich damit zu einer extrem harten Kugel zusammenziehen können.
Alle C. acus wurden zusammen mit Seewalzen in Meerwasseraquarien gehalten. Die
Versuche wurden in einem Glasaquarium (40 x 20 x 25 cm) und in einem Plastik-
aquarıum (50 x 30 x 30 cm) durchgeführt. Um es den wegen der Fangbedingungen
durch Dekompressionsschäden geschwächten Fischen zu ermöglichen in die Seewalze
BIOLOGIE VON “FIERASFER” (CARAPUS ACUS) 337
zu gelangen, wurde der Wasserspiegel während der ersten Tage nach dem Fang bis auf
Höhe des Afters der Seewalze gesenkt. Für die Beobachtung des Eindringens wurde
jeweils ein C. acus zusammen mit einer H. tubulosa oder einem S. regalis in ein
Aquarium gesetzt. Bei den Wahlversuchen bekam der Fisch immer eine H. tubulosa
und einen S. regalis, die wahllos aus der Menge der vorhandenen Seewalzen heraus-
gegriffen worden waren. Wenn C. acus in einer Seewalze verschwunden war, wurde
das Verhalten des Wirtes mindestens 5 min weiter beobachtet, um zu prüfen ob der
Fisch ihn wieder verläßt. Nach Versuchsende wurde C. acus mit der Seewalze wieder
in das Hälterungsbecken zurückgesetzt. Die Dokumentation der Versuche erfolgte
exemplarisch mittels Videokamera (Typ: Sony Video 8) und Photoapparat (Typ: Canon
EOS 100). Ansonsten wurden schriftliche Protokolle erstellt. Um zu klären welche
Faktoren für die Art des Einstiegs in die Seewalze verantwortlich sind, wurde das vor-
und rückwärtige Eindringen der Fische in Einzelbildsequenzen aufgelöst. Zur
Auswertung dieser Sequenzen dienten ein Videorecorder (Typ: Panasonic) und ein
Computer (Programm: Scancam).
Auch alle für die anatomischen und histologischen Untersuchungen verwen-
deten C. acus Exemplare entstammten S. regalis aus 70 - 110 m Meerestiefe. Die
Fische wurden in 10% Formalin oder nach Bouin fixiert und später in 70% Ethyl-
alkohol überführt. Sie wurden anschließend vermessen, der Länge nach sortiert,
numeriert und ihr Geschlecht anhand der Gonaden bestimmt. Im Labor wurden die
Mägen von in Formalin fixierten C. acus unter dem Binokular geöffnet und auf ihren
Inhalt untersucht. Die dabei gefundenen Kleinkrebse wurden mit Boraxkarmin gefärbt
und in Kanadabalsam eingeschlossen. Bei nach Bouin fixierten Fischen wurden die
Magen ganz herauspräpariert und in Paraffin eingebettet. Von den Paraffinblöcken
wurden 7um dicke Schnitte angefertigt, mit Hämalaun-Eosin oder Azan gefärbt und
unter einem Zeiss-Mikroskop ausgewertet und photographiert.
ERGEBNISSE
Die Häufigkeit von C. acus in S. regalis war in den verschiedenen Jahren sehr
unterschiedlich: während 1995 im Juli 25 C. acus in 45 S. regalis gefunden wurden;
waren es 1998 zur selben Jahreszeit nur 12 Fische in 329 S. regalis; 1996, 1997 und
1999 fischten wir keinen einzigen C. acus. Der Besatz von S. regalis mit C. acus war
1995 mit 55% viel höher als in allen anderen Jahren, obwohl alle Fänge zur gleichen
Jahres- und Tageszeit im selben Gebiet stattfanden und auch die angewandten Fang-
methoden, sowie Boot, Netz und Mannschaft immer die gleichen waren. Die beiden
einzigen C. acus aus 85 H. tubulosa wurden 1998 in weniger als 10 m Meerestiefe über
Felsgrund bei Banyuls-sur-mer gefunden.
Die Stichopus-Exemplare waren aus 70 - 110 m Tiefe geholt worden, weshalb
alle enthaltenen Fische deutliche Dekompressionsschäden zeigten, die 3 - 4 Stunden
nachdem sie auf Meereshöhe gebracht worden waren in Form von Gasbläschen auf-
traten. Im Bereich von Kopf und Magen bildeten sich Gasansammlungen, die manch-
mal zusätzlich über den ganzen Schwanz verteilt vorkamen. Weil der Auftrieb der
Fische durch diese im Körper eingelagerten Gase erheblich erhöht wurde, war es ihnen
338 KERSTIN KLOSS UND WOLFGANG PFEIFFER
unmöglich im Aquarium zu den Seewalzen am Grund abzutauchen. Zudem wirkten die
Fische extrem geschwächt. Nur in der Seewalze erholten sie sich; alle C. acus, die es
nicht schafften sich in eine Seewalze zurückzuziehen, erlagen innerhalb weniger Tage
ihren Verletzungen. Andere Exemplare mit ähnlich schwerwiegenden Dekompressions-
schäden erholten sich in der Seewalze binnen maximal 3 Tagen und konnten danach
auch bei normalem Wasserstand wieder zum After abtauchen um in die Seewalze
einzudringen.
Wahlversuche: Da die meisten Individuen von C. acus in S. regalis gefunden
worden waren, sollte überprüft werden ob S. regalis auch bei einer Wahlmöglichkeit
bevorzugt wird. Alle Wahlversuche verliefen überraschend gleichförmig. C. acus zeigte
immer ein ausgesprochenes Interesse an A. tubulosa, was am schnelleren und weiter
ausholenden Schlängeln seines Schwanzes zu erkennen war. Stieß der Fisch zufällig
zuerst auf S. regalis, machte er keinerlei Anstalten in sie einzudringen, sondern suchte
das Aquarium weiter ab. Traf er zuerst auf A. tubulosa, ging er meist sofort in “Lauer-
stellung”. Diese Stellung ist dadurch gekennzeichnet, daß Kopf und Rumpf regungslos
bleiben während der Schwanz eine schnelle Schlängelbewegung durchführt. Der
Körper nimmt dabei vor dem After der Seewalze eine horizontale Lage ein. Aus dieser
Lauerstellung heraus versucht C. acus in den After der Seewalze zu gelangen sobald
diese Atemwasser ausstößt. In insgesamt 11 Wahlversuchen mit 3 Fischen drangen alle
ohne Ausnahme in A. tubulosa ein: Fisch Nr. 30 siebenmal, Nr. B und Nr. 31 je
zweimal. In den Versuchen mit den Fischen Nr. B und Nr. 31 wurde jeweils einmal
beobachtet, wie sie in eine schon besetzte A. tubulosa eindrangen, obwohl jedesmal ein
unbewohnter S. regalis als Alternative zur Verfügung stand. Auch in anderen Ver-
suchen hielten sich wiederholt 2 - 3 C. acus in einer A. tubulosa oder im selben
S. regalis auf. Die Wahlversuche zeigten deutlich, daß C. acus H. tubulosa gegenüber
S. regalis bevorzugt.
Einstiegsvarianten: C. acus kann auf 2 verschiedene Weisen in die Seewalze
eindringen - Kopf oder Schwanz voraus. Kopf voraus Eindringen (Abb. 1): C. acus legt
sich in Lauerstellung vor den After der Seewalze. Sobald sie ihren Atemwasserstrom
durch den After ausstößt, beschleunigt der Fisch blitzschnell die Schlängelbewegungen
seines Schwanzes und ist innerhalb von 40-80 ms vollständig in der Seewalze
verschwunden. Dieser Vorgang ist manchmal so schnell, daß die Einzelbildanalyse
keinen genauen Aufschlufl ermöglicht: bei S. regalis: oft schneller als 80 ms, manchmal
aber auch 4, 10, 13, 50 und 65 s; bei H. tubulosa meist zu schnell um mit Aufnahmen
von 25 Bildern/ s festgehalten werden zu können, oft 40-80 ms, nur ausnahmsweise
mehrere Sekunden. Schwanz voraus Eindringen (Abb. 2a-c): Wenn C. acus rückwärts
in die Seewalze einzudringen versucht, nimmt er zuerst dieselbe Lauerstellung ein wie
beim Vorwärtseindringen. Jetzt ist aber zu beobachten, daß die Schwanzspitze sich
immer wieder an der rechten oder linken Seite einzurollen und eine Schleife zu bilden
beginnt. Spürt der Fisch den Wasserstrom der Seewalze, führt er den Schwanz an seiner
Körperseite und am Kopf vorbei, den Körper gleichsam als Gleitschiene verwendend.
Der Kopf dient dabei als Einfädelhilfe für den Schwanz, so daß dieser in den geöffneten
Anus eingeführt werden kann. Die Fische bewegten hierbei den Schwanz in fast allen
Fällen an ihrer linken Körperseite vorbei. Steckt der Fisch bereits mit der
BIOLOGIE VON “FIERASFER” (CARAPUS ACUS) 339
ABB. 1. Carapus acus (Länge 115 mm): Vorwärtseindringen in S. regalis; a) Lauerstellung mit
Maul am After der Seewalze; b) Kopf in Seewalze, kräftiger Schwanzschlag. (Fotos: T. Moritz)
340 KERSTIN KLOSS UND WOLFGANG PFEIFFER
Schwanzspitze im After der Seewalze, so biegt er seinen Körper nach einer Seite hoch,
wodurch ihm anscheinend ein besserer Halt ermöglicht wird. Im Innern der Seewalze
verkeilt er sich. Je nach Seewalzenart und Fischgröße dauerte es 12 - 472 s bis C. acus
ganz in der Seewalze verschwunden war: bei S. regalis: 15, 34, 58, 102, 139 und 195 s;
bei A. tubulosa: 12, 13, 32, 93, 110 und 472 s. Der Rückwärtseintritt währte mehr als
doppelt so lang wie der Vorwärtseintritt. Unterbrechungen im Ablauf des Eindringens
fanden an unterschiedlichen Stellen statt. Daß der Fisch dabei durch seine leicht
abstehenden Brustflossen und Kiemendeckel behindert wird, war nicht zu erkennen.
Nachdem C. acus ganz in der Seewalze verschwunden war, wurde wiederholt
beobachtet wie er seinen Kopf durch den After der Seewalze teilweise herausstreckte
und atmete (Abb. 2d). In manchen Fällen besaßen S. regalis-Exemplare, in welche
Fische eindrangen, keine weiteren inneren Organe mehr außer Darmresten. Gonade und
Wasserlunge waren bereits früher ausgeworfen worden. In diesen Fällen ist anzu-
nehmen, daß der Fisch über dieselbe Öffnung der Enddarmerweiterung in die Leibes-
höhle vordrang über die er in die Wasserlunge gelangen könnte. Problematisch wird die
Erklärung allerdings für Fische, die in der Leibeshöhle von Seewalzen gefunden
wurden, welche noch über ihre Wasserlunge verfügten. Es wird vermutet, daß sich der
Fisch in der engen Wasserlunge umdreht und dabei das dünnhäutige Lungengewebe
durchstößt.
Drei S. regalis wurden beobachtet wie sie auf das Eindringen des Fisches
umgehend mit heftigem Schlängeln reagierten. Ihre Auf- und Abbewegungen dauerten
bis zu 8 min und wurden auch durch Umsetzen in ein anderes Aquarium nicht
unterbrochen; der C. acus im Innern verließ seine Behausung jedoch nicht. A. tubulosa
versuchte durch Kontraktion ihres Afterschließmuskels und ihrer Hautmuskulatur im
hinteren Körperabschnitt den Fisch am Eindringen zu hindern. Ihr Körper erschien
dadurch im posterioren Bereich wesentlich dünner. Wie die lange Einstiegsdauer
vermuten läßt, wurde dadurch das Vorhaben von C. acus zwar erschwert, doch nicht
verhindert. In 2 Fällen kotete H. tubulosa, wodurch es ihr gelang den bereits teilweise
eingedrungenen C. acus wieder herauszudrücken. Beide Male startete der Fisch
sogleich einen weiteren Versuch, der jeweils schnell zum Erfolg führte. Häufigkeit und
Verteilung der vor- und rückwärtigen Eintritte von C. acus hängen sowohl von der
Größe des Fisches als auch von der Seewalzen-Species ab. Bei Fisch Nr. 30 (115 mm)
halten sich die rück- und vorwärtigen Eintritte in S. regalis mit 9:7 etwa die Waage; bei
H. tubulosa wurde dagegen der Rückwärtseinstieg mit 20:5 bevorzugt. Zwei weitere
Fische entschieden sich mit 11:2 bzw. 7:0 ebenfalls für den Riickwartseintritt. Der
Wechsel zwischen den beiden Eintrittsvarianten erfolgte bei diesen Fischen mittlerer
Körperlänge unregelmäßig. Größere Fische drangen dagegen immer mit dem Schwanz
voran ein, egal ob sie S. regalis oder H. tubulosa vor sich hatten. Haut- und Schließ-
muskulatur sind bei H. tubulosa stärker entwickelt und kräftiger als bei S. regalis, bei
dem sich der After auch in kontrahiertem Zustand leicht öffnen läßt. Der Fisch kann
also auch mit seinem dicken Vorderende voraus eindringen. Anders ist die Situation bei
H. tubulosa. Mit ihrer kräftigen Muskulatur verschließt sie den Anus so fest, daß es
selbst mit einer dünnen Sonde nahezu unmöglich ist ihn zu öffnen. Hier steckt C. acus
BIOLOGIE VON “FIERASFER” (CARAPUS ACUS) 341
ABB. 2. Carapus acus (dasselbe Exemplar wie in Abb. 1): (a-c) Rückwärtseindringen in A.
tubulosa; a) Einrollen der Schwanzspitze; b) Schwanz wird links am Körper vorbeigeführt; c)
Fisch hat sich umgedreht, Schwanzspitze im After der Seewalze; d) Kopf von oben, in S. regalis;
e) beim Freipräparieren in S. regalis. (Fotos: T. Moritz)
342 KERSTIN KLOSS UND WOLFGANG PFEIFFER
zuerst seinen spitz auslaufenden Schwanz als dünneres Körperende in den Anus. Ist der
Schwanz erst einmal in der Seewalze, kann den Fisch fast nichts mehr davon abhalten
ganz einzudringen. Handelt es sich bei dem Fisch um ein größeres Exemplar,
verwendet er die Variante “Schwanz voraus” auch bei S. regalis, da sein Kopf für einen
frontalen Vorstoß zu dick ist.
Mageninhalt: In histologischen Schnittpräparaten von 2 nach Bouin fixierten
Mägen sind deutlich Komplexaugen von Kleinkrebsen zu erkennen. Dagegen konnten
histologisch keine Hinweise darauf gefunden werden, daß C. acus auch Gewebe der
Seewalzen fressen würde. Weder Gonaden noch Muskulatur oder sonstige zelluläre
Strukturen von S. regalis befanden sich in den Fischmägen. In den Magen von 2 in
Formalin fixierten Fischen aus S. regalis, wurden 4 Kleinkrebse angetroffen bei denen
es sich vermutlich um die Garnele Pandalina sp. (Decapoda) handelt (Fischer er al.,
1987). Insgesamt wurden in den Mägen von 4 der 17 untersuchten Männchen und
Weibchen von C. acus (154 - 195 mm Gesamtlänge) Krebse oder deren Augen
gefunden. Knapp 1/4 der Fische hatte also kurz vor seinem Fang Krebse gefressen.
DISKUSSION
VERBREITUNG: Die meisten Species der Gattung Carapus leben in tropischen
oder subtropischen, überwiegend seichten Gewässern (Williams, 1984; Markle &
Olney, 1990). Carapus acus ist auf das Mittelmeer und die angrenzende südliche
Atlantikküste beschränkt. Arnold (1953, 1956) fand C. acus bei Neapel hauptsächlich
in H. tubulosa aus 10 - 20 m Meerestiefe. Von 1350 bei Neapel in 10 - 25 m Tiefe
gesammelten A. tubulosa-Exemplaren enthielten nur 19 (1,4%) einen C. acus (Gustato,
1976). Der Befall von nur 2 A. tubulosa bei Banyuls-sur-mer hatte den Anschein von
Zufälligkeit, denn 38 von 40 C. acus fanden wir in S. regalis aus 70 - 110 m Tiefe. Die
im Juli - August 1998 gefangenen C. acus waren überwiegend Jungfische (Gesamtlänge
durchschnittlich 129 mm), von denen sich aber keiner mehr in einem Larvenstadium
befand. Im Gegensatz dazu handelt es sich bei den 1995 zur selben Jahreszeit gefan-
genen Fischen hauptsächlich um große adulte Exemplare (Gesamtlänge durchschnitt-
lich 173 mm), wobei der Befall 55% betrug. Im Meer haben wir zwar die meisten (38)
C. acus in (450) S. regalis und nur 2 in (85) H. tubulosa gefunden, doch als wir die
Fische im Labor vor die Wahl zwischen S. regalis und H. tubulosa stellten, entschied
sich C. acus immer für H. tubulosa. Die Bevorzugung von A. tubulosa im Aquarium
war überraschend, da deren Tiefenverbreitung im Untersuchungsgebiet von Banyuls-
sur-mer großteils nicht mit der des Fisches übereinstimmt. Bei den zwei in geringer
Tiefe in A. tubulosa gefangenen C. acus handelte es sich vermutlich um verirrte Indivi-
duen, obgleich C. acus in anderen Regionen vor allem in A. tubulosa aus geringer Tiefe
gefischt wurde (Emery, 1880). Der Vergleich der Literaturangaben zur Tiefenver-
breitung von C. acus zeigt hingegen, daß sich das Vorkommen des Fisches mit dem
beider Seewalzen-Species in anderen Gebieten überschneidet (Abb. 3). Folgende Hypo-
these könnte die experimentelle Bevorzugung von H. tubulosa erklären. Geht man
davon aus, daß sich der Fisch von Kleinkrebsen ernährt, wäre es gleichgültig welche
Species er als “Wohnhöhle” aussucht. Wichtig wäre dagegen, welches Substrat die See-
|
|
BIOLOGIE VON “FIERASFER” (CARAPUS ACUS) 343
walze bevorzugt, da auch die Beutekrebse untergrundspecifische Vorlieben haben. Die
als Nahrung nachgewiesene Pandalina sp. lebt überwiegend auf Felsgrund. Auch A.
tubulosa wurde meistens in Felsnähe gefunden. Als Substratfresser ist diese Seewalze
zwar auf Sandgrund angewiesen, lebt jedoch vor Banyuls-sur-mer hauptsächlich an
Orten, an denen die schützende Felsformation der Küste in sandigen Untergrund
übergeht. Vor Banyuls-sur-mer ist dies in 5 - 20 m Tiefe der Fall. S. regalis kommt
hingegen vorwiegend auf Sandgrund vor. So ließe sich erklären, warum C. acus H.
tubulosa gegenüber S. regalis bevorzugt. In der nur bis ca. 25 m tiefen Bucht von Port
de la Selva mit Sandgrund und Posidonia-Wiesen sind H. tubulosa und besonders
H. polii zwar häufig, doch fehlen hier C. acus und S. regalis.
VERHALTEN: Aus einer Seewalze herauspräparierte und im Aquarium frei
schwimmende C. acus zeigten zwei völlig unterschiedliche Schwimmweisen. Die einen
schwammen an der Wasseroberfläche mit weit ausholenden schlängelnden Bewegun-
gen und streckten dabei ihren Kopf immer wieder aus dem Wasser. Die anderen
tauchten sofort Kopf voran zum Aquarienboden, wobei ihre schräge Schwimmlage
beibehalten wurde bis sie auf eine Seewalze stießen. Arnold (1953) beschreibt ebenfalls
diese beiden Schwimmweisen, deutet sie jedoch als Kennzeichen verschiedener Ent-
wicklungsstadien. Nach Arnold (1953) schwimmen Tenuis-Larven mit weitausholen-
den Bewegungen während junge und adulte Fische nur leicht mit dem Schwanz
schlängeln. Unsere Beobachtungen konnten diese Aussage nicht bestätigen. Vielmehr
hatte es den Anschein, daß die Schwimmweise mit dem Allgemeinbefinden des Fisches
zusammenhängt. Hatte er Dekompressionsschäden oder war der Sauerstoffgehalt im
Aquarium zu niedrig, so schnappte er wiederholt an der Oberfläche nach Luft und
schwamm mit hektischen Bewegungen. Das gleiche Verhalten zeigte er auch in
Bedrängnis. Dagegen schwamm er bei gutem Allgemeinzustand in Schräglage mit dem
Maul am Boden. Die beiden Schwimmweisen sind somit nicht abhängig vom
Entwicklungsstadium sondern Ausdruck des Befindens. Wenn der Fisch vor dem After
der Seewalze seine Lauerstellung einnimmt, sind die Bewegungen der Brustflossen
deutlich zu erkennen. Diese Flossen schlagen abwechselnd und dienen sowohl der
Balance als in geringem Maße auch der Lokomotion. Der Vortrieb erfolgt haupt-
sächlich über den Schwanz, wie beim Vorwärtseindringen zu sehen ist. Hierbei
beschleunigt der Fisch in kürzester Zeit bis auf Höchstgeschwindigkeit durch kräftiges,
weitausholendes Schwanzschlagen.
Carapus acus findet die Seewalzen vermutlich zufällig; ein besonderes Such-
schema war nicht zu erkennen. Nach Arnold (1958) und Trott (1981) geschehe die
Lokalisierung von weiter entfernten Zielen chemisch, während auf kurze Distanz die
visuelle Orientierung eine Rolle spiele. Van Meter & Ache (1974) vermuten, daß nur
die Lokalisierung des Afters über die Augen erfolgt. Unsere Fische schwammen im
Aquarium umher bis sie auf eine Seewalze trafen. Ob und wie weit hierbei chemische,
taktile oder visuelle Reize eine Rolle spielen, kann nicht entschieden werden. Die
deutliche Reaktion auf das ausgestoßene Atemwasser ist wahrscheinlich taktil und
chemisch bedingt. Eine visuelle Wahrnehmung ist nicht anzunehmen, da fast alle
Versuchsfische aus 70 - 110 m Tiefe stammen und ihre Augen weitgehend rückgebildet
sind. Die Beobachtung, daß vor allem kleinere Fische vorwärts in die Seewalze
344 KERSTIN KLOSS UND WOLFGANG PFEIFFER
eindringen, bestätigt Trott’s (1981) Aussage, daß diese Weise die schnellste Möglich-
keit des Eindringens darstellt. Die Entscheidung, ob ein Fisch vorwärts oder rückwärts
eindringt, wird durch die morphologischen Gegebenheiten der Seewalzen-Species und
die Fischgröße bestimmt, denn auch große Fische können vorwärts und kleine rück-
warts eindringen. Zweimal wurden am Morgen Fische freischwimmend im Aquarium
angetroffen, die sich am Vorabend noch in einer Seewalze befunden hatten. Ferner
entwischten einige Exemplare in der Nacht durch die unzureichend gesicherten über-
läufe. Dies zeigt, daß C. acus die Seewalze nachts verläßt, vermutlich um auf Beute-
fang zu gehen. Einmal wurde beobachtet, wie ein C. acus seine H. tubulosa verließ
nachdem er etwa eine halbe Stunde in ihr verweilt hatte. Der Fisch schwamm einige
Runden im Aquarıum und verschwand dann wieder in derselben Seewalze. Das
selbständige Verlassen der Seewalze, ohne von ihr ausgeworfen zu werden, ist also
möglich. Daß sich C. acus nicht dauernd in der Seewalze aufhält, zeigen auch die im
Meer freischwimmenden Exemplare (Smith, 1964).
ERNÄHRUNG: Die anatomischen und histologischen Untersuchungen bewiesen,
daß C. acus Krebse frißt. Fraglich blieb hierbei, ob diese sein ganzes Nahrungs-
spektrum ausmachen. Es gibt jedoch keinen Hinweis, daß C. acus seinen Wirt anfrißt.
In keinem der untersuchten Mägen wurden Reste von S. regalis gefunden. Diese
Befunde sprechen für die Annahme, daß C. acus kein Nahrungsparasit ist. Während der
Verhaltensexperimente hatte C. acus nur Seewalzen zur Verfügung von denen er sich
laut Arnold (1953) parasitisch ernährt. Eine andere potentielle Nahrung wurde in
unseren Wahlexperimenten nicht angeboten. Auf separate Fütterungsversuche mit
Artemia sp. oder lebenden Larven von Mysis sp. reagierte C. acus nicht. Wurden die
Futtertiere ins Aquarium gebracht, so schwamm der Fisch sofort in die dunkelste Ecke
ohne sich um die Krebse zu kümmern. Pandalina sp., die wir in den Magen von C. acus
gefunden hatten, war für uns als Futtertier nicht zugänglich. Gab man die Wasserlunge
von S. regalis oder H. tubulosa ins Aquarium, war wiederholt zu beobachten, daß sich
der Fisch neben oder unter der Lunge zusammenringelte. Kurzzeitig hatten wir deshalb
die Coelomocyten der Seewalzen, die auch der Verdauung und Nährstoffspeicherung
dienen, in Verdacht von C. acus aufgenommen zu werden. In diesem Fall müßten sich
die Fische trinkend ernähren, indem sie die Coelomflüssigkeit der Seewalze schlucken
um die in den Coelomocyten gespeicherten Stoffe zu verwerten. Zum einen besitzt
C. acus keine Strukturen die einen filtrierenden Nahrungserwerb ermöglichen, zum
anderen müflten gewaltige Mengen von Coelomocyten geschluckt werden um den
Energiebedarf zu decken.
Um die Frage eines “Zubrotes” neben Futterkrebsen zu klären, wurde zum
Vergleich die freilebende Carapinae-Art Echiodon drummondi Thompson, 1837 un-
tersucht (Trott, 1981). Diese lebt in größeren Tiefen (bis ca. 300 m) vor den Küsten
Britanniens, des südlichen Norwegens, des westlichen Dänemarks und in der Biskaya
(Trott & Olney, 1986); sie ernährt sich von kleinen Invertebraten und Fischen.
Achtzehn Exemplare von E. drummondi, die der Zweit-Autor 1985 vor Arcachon in der
Biskaya in 200-300 m Meerestiefe gefangen und in Formalin fixiert hatte, wurden auf
ihren Mageninhalt geprüft. In einem wurde ein vermutlich parasitischer Nematode
gefunden, 6 andere enthielten Krebse. Wegen der fortgeschrittenen Verdauung waren 3
BIOLOGIE VON “FIERASFER” (CARAPUS ACUS) 345
100
120
400
AO) FC) 20) Se) eo) mel. 0)
We bis unterhalb von [ | Hauptverbreitung
IN bis oberhalb von | È seltenes Vorkommen
ABB. 3. Vertikale Verbreitung von S. regalis, H. tubulosa und Carapus acus: Vergleich der
eigenen Befunde (schwarz) mit Literaturangaben (weiß) von: a) Nadal i Fortia (1981, 1994), b)
Fiedler & Lieder (1994), c) Riedl (1983), d) Mayer (1937), e) Bauchot & Pras (1980), f) Fiedler
(1991), g) Markle & Olney (1990). Ordinate: Tiefe in m.
Krebse nicht zu identifizieren, bei den 3 anderen handelte es sich um Isopoda.
Insgesamt hatte also 1/3 der untersuchten Fische Crustaceen im Magen. Bei C. acus
betrug der Anteil der Exemplare mit Krebsen im Magen etwa 1/4. Der Vergleich der
beiden Werte legt die Vermutung nahe, daß auch C. acus kein Nahrungsparasit ist,
sondern ein nachtaktiver Jäger. Es ist anzunehmen, daß die tagsüber gefischten C. acus
ihr nächtliches Mahl bereits verdaut hatten. Demgegenüber müßten bei parasitischer
346 KERSTIN KLOSS UND WOLFGANG PFEIFFER
Ernährung, die keinem tageszeitlichen Faktor unterliegt, ständig Gonaden oder andere
Gewebe von Seewalzen in den Fischmägen gefunden werden. Dies war in unserer
Untersuchung jedoch nicht der Fall. Arnold (1953), der die parasitische Ernährung für
C. acus postuliert, führt keinen Beweis für seine Spekulation an. Außerdem beobachtete
Gustato (1976), daß von 17 C. acus, die er aus H. tubulosa entnahm, sich nur 2 in der
Leibeshöhle befanden. Wie sollten die restlichen 15 Fische in der Wasserlunge von der
Gonade fressen?
Um die Hypothese zu untermauern, wonach sich C. acus rein räuberisch ernährt,
ist ein Vergleich mit den anderen Species der Gattung Carapus hilfreich. Für viele von
ihnen wird ebenfalls eine räuberische Lebensweise angenommen. Krebse wurden
gefunden im Magen von C. bermudensis (Smith et al., 1981), C. dubius (Trott, 1970),
C. homei (Smith, 1964) und C. mourlani (Trott, 1970). Auch C. birpex (Trott & Olney,
1986) und C. parvipinnis (Fiedler, 1991) werden nicht Parasiten genannt, sondern wie
die vier vorher aufgelisteten Species als Kommensalen, Einmieter, Inquilinisten,
Kannibalen oder Symbionten bezeichnet (Strasburg, 1961, Smith & Tyler, 1969;
Seymour & Mc Cosker, 1970; Dawson, 1971, van Meter & Ache, 1974; Markle &
Olney, 1980; Fiedler, 1981; Trott, 1981; Trott & Olney, 1986).
All diese Befunde stützen die Hypothese, wonach auch C. acus als Kommensale
lebt. Als bessere ökologische Begriffe für die Lebensweise von C. acus können
Endophoresie (Baer, 1951) und Entökie (Hohorst, 1981; Hentschel & Wagner, 1984)
herangezogen werden. Am treffendsten erscheint uns die Bezeichnung Inquilinismus
(Emery, 1880; Bertin & Arambourg, 1958; Trott, 1970; Trott & Olney, 1986), da sich
der Fisch selbständig ernährt. Völlig fehl am Platz ist hingegen der Begriff Symbiose
(van Meter & Ache, 1974; Gustato, 1976), da aus unserer Sicht die als Herberge
dienende Seewalze keinerlei Vorteil von ihrem Mieter hat, der anscheinend allein
profitiert. Aufgrund Arnold’s (1953) unbewiesener Vermutung ist C. acus bisher zu
Unrecht als Nahrungsparasit bezeichnet worden. Diese Behauptung wurde von der
umfangreichen Sekundärliteratur übernommen (Bauchot & Pras, 1980; Riedl, 1983;
Terofal, 1986; Fiedler, 1991; Fiedler & Lieder, 1994 u.a.), was zur heutigen all-
gemeinen Verbreitung dieser Ansicht führte. Es gibt jedoch bisher keinen einzigen
Beweis dafür, daß C. acus von der Seewalze frißt und sich somit parasitisch ernährt.
Die Seewalze dient ihm vielmehr nur als schützende Zuflucht zwischen seinen nacht-
lichen Beutezügen. Carapus acus muß dementsprechend als Inquilinist bezeichnet
werden, wie dies Emery (1880) bereits völlig richtig getan hat - er nennt den Fisch
“inquilino”!
SUMMARY
450 Stichopus regalis (Cuvier), caught with a trawl-net at a depth of 70 - 110 m
off Banyuls-sur-mer (France), contained 38 pearlfish, Carapus acus (Brünnich). Two
pearlfish were found in 85 Holothuria tubulosa Gmelin caught at Banyuls-sur-mer in
less than 10 m of seawater. In contrast, 63 H. tubulosa and 142 H. polii Della Chiaje,
collected by diving in the bay of Port de la Selva (ca. 10 miles south of Banyuls), did
not contain any C. acus. Pearlfish collected from 1995 to 1998 were preserved to study
BIOLOGIE VON “FIERASFER” (CARAPUS ACUS) 347
their stomach contents. Twelve individuals sampled in the summer of 1998 were kept in
aquaria to analyse their behaviour by means of computer analysis of video records. Due
to severe damage caused by insufficient decompression only a few individuals survived
longer than one week after being placed in an aquarium. Since most C. acus were
collected from S. regalis, the host preference of the fish was tested in an aquarium.
S. regalis and H. tubulosa were provided as potential hosts. The results were surprising:
in 11 experiments the 3 fish examined entered only H. tubulosa if they could choose
between both species.
The entry of the pearlfish through the anus of the host holothurian may be head
first or tail first. The pearlfish, in a head-down position, lies in wait in front of a sea
cucumber. As the sea cucumber discharges its respiratory water the fish rapidly flicks
its tail and enters the host. In the case of H. tubulosa the pearlfish enters head first
within 40 - 80 ms; in S. regalis it takes up to 4 - 60 s. Tail first entry of C. acus takes 12
- 120 s. To enter the host, the pearlfish bends up its tail towards the head and its whole
body forms a loop. When the waiting pearlfish perceives the water discharged by a
holothurian, it moves its tail foreward along the side of its body and head. By this, the
head helps the tip of the tail to target the anus of the host. Three S. regalis responded to
intruding pearlfish with intense up and down movements; H. tubulosa contracted the
sphincter of its anus and the spherical muscles of its body wall in the posterior region,
whereby they became thin. However, neither defence method prevented C. acus from
entering its host. In C. acus the frequency of head first and tail first entries depended on
the size of the pearlfish and on the species of the host holothurian. A 145 mm long
pearlfish entered S. regalis 9 times tail first and 7 times head first. However, the same
fish entered H. tubulosa 20 times tail first and only 5 times head first.
The presence of C. acus in S. regalis sampled was extremely variable. In 1995
as many as 25 C. acus were collected from 45 S. regalis specimens. However, since
1996 only 13 C. acus were found in 405 S. regalis, although the locality, depth, season,
day time, boat, trawl-net, sampling method and techniques, and even the crew were the
same. The C. acus specimens sampled in 1995 were large (0 173 mm) and adult,
whereas the pearlfish caught in 1998 were much smaller (@ 129 mm) and mostly
juvenile.
Pearlfish removed from the host holothurian demonstrated two completely
different styles of swimming. Either they swam at the water surface with wriggling
movements, repeatedly raising their head out of the water, or they immediately dived to
the ground of the tank, maintaining a 30 - 40 degree tilted head-down position until
they found a host holothurian. The latter style of swimming is only exhibited by healthy
individuals. Swimming at the surface indicates either damage caused by insufficient
decompression or a poor physical condition of the pearlfish.
In the aquarium it was found that pearlfish occasionally leave their host holo-
thurians, especially at night, when they presumably feed. The stomachs of 4 out of 17
individuals contained small shrimps, probably Pandalina sp. In comparison, isopods
and other small crustaceans were found in the stomachs of 6 out of 18 specimens of the
closely related, but free-living, Echiodon drummondi Thompson. The statement by
Arnold (1953) that C. acus is a parasite which feeds on the organs (gonads etc.) of its
348 KERSTIN KLOSS UND WOLFGANG PFEIFFER
sea cucumber host could not be confirmed. This statement is completely speculative
and not based on any research results. The stomachs of C. acus examined did not
contain any holothurian tissue. All other species of the genus Carapus feed on small
invertebrates and fish, and none feed parasitic. Our results give reason to agree with
Emery (1880), who stated that C. acus is an inquiline (“inquilino”), to whom the host
holothurian serves as a shelter only.
DANKSAGUNG
Wir danken all denen, die geholfen haben, daß diese Arbeit zustande kam. Unser
besonderer Dank gebührt der Direktion des Laboratoire Arago in Banyuls-sur-mer
(Frankreich), die uns ermöglichte, auf diesem Gebiet zu arbeiten. Die Mitarbeiter der
Station waren immer hilfsbereit. Weder schlechtes Wetter noch stürmische See konnten
die Mannschaft des Forschungsschiffes Nereis davon abhalten mit uns auszufahren, um
Seewalzen zu fischen. Besonders zu erwähnen ist unser Kollege Timo Moritz, der
große Geduld bewies beim photographischen Festhalten des Eindringens von C. acus in
Seewalzen.
LITERATUR
ARNOLD, D. C. 1953. Observations on Carapus acus, (Brunnich) (Jugulares, Carapidae). Pubbli-
cazioni della Stazione Zoologica di Napoli 24: 152-166.
ARNOLD, D. C. 1956. A systematic revision of the fishes of the Teleost family Carapidae (Perco-
morphi, Blennioidea), with descriptions of two new species. The Bulletin of the British
Museum (Natural History) 4: 247-307.
ARNOLD, D. C. 1958. Further studies on the behaviour of the fish Carapus acus (Briinnich).
Pubblicazioni della Stazione Zoologica di Napoli 30: 263-268.
BAER, J. G. 1951. Ecology of animal parasites (pp. 1-7). The University of Illinois Press, Urbana,
224 pp.
BAUCHOT, M. L. & Pras, A. 1980. Guide des poissons marins d’ Europe, pp.196-197. Delachaux
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REVUE SUISSE DE ZOOLOGIE, 107 (2): 351-357; juin 2000
Un nouveau Collembole Neanurinae du Sud du Vietnam,
Blasconura batäi sp. n., avec une clé des especes du genre
(Collembola: Neanuridae)
Anne BEDOS & Louis DEHARVENG
Laboratoire d’Ecologie Terrestre, UMR 5552 du CNRS,
U.P.S., 118 route de Narbonne, 31062 Toulouse Cedex 4, France.
E-mail: deharven @cict.fr
A new neanurine springtail from southern Vietnam, Blasconura batai
sp. n., with a key to the species of the genus (Collembola: Neanuridae).
- The new species is described from a karstic area in southern Vietnam. A
checklist and a key of all species of the genus Blasconura are given.
Key-words: Collembola - Neanuridae - Vietnam.
INTRODUCTION
Les collines de Hon Chong, isolées en bordure du Golfe de Thailande dans le
Sud du Vietnam, constituent un remarquable site karstique dont une grande partie a été
récemment détruite par l’extension de carrieres pour l’exploitation du calcaire. Nous
avons pu y réaliser quelques récoltes qui nous ont permis de décrire deux especes de
Collemboles, Lepidonella lecongkieti Deharveng & Bedos, 1995 et Rambutsinella
honchongensis Deharveng & Bedos, 1996. Nous décrivons dans le présent travail une
troisieme espece de cette région, Blasconura batai sp. n., et proposons une clé des
especes du genre Blasconura.
Blasconura Cassagnau, 1983
Le genre Blasconura a été créé pour l’espece Achorutes hirtellus Boerner, 1906
de Java. La description originale de cette espèce, très sommaire, peut s’appliquer à de
nombreux Neanurinae tropicaux de la tribu des Paleonurini. Il n’est donc pas étonnant,
étant donné la confusion systématique qui régnait au sein des Neanurinae avant les
mises au point de Cassagnau (1983, 1986, 1989), que l’espèce hirtella ait été signalée
dans de nombreuses régions d’Asie du Sud-Est jusqu’en Australasie. Cette vaste distri-
bution concerne probablement des espèces différentes, comme en témoignent les illus-
trations données dans la littérature (Handschin, 1926; Womersley, 1933, 1936; Denis,
Manuscrit accepté le 18.11.1999
32 ANNE BEDOS & LOUIS DEHARVENG
1948; Yosu, 1959, 1966; Gapud, 1968). Aucune de ces formes ne peut être attribuée
sans ambiguïté au véritable B. hirtella tel que l’a redécrit Yosii (1976) d’après un
matériel topotypique de Bogor (Java), ce que confirment nos propres observations sur
de nouveaux exemplaires provenant de la localité-type. Autrement dit, toutes les
citations de B. hirtella antérieures au travail de Yosu (1976) devront être révisées.
En 1988, Cassagnau décrit 6 Blasconura nouveaux d’Inde et de Sri Lanka; il
suggere alors de placer Achorutes separatus Denis, 1934 du Vietnam dans le genre
Blasconura. En suivant cette conception, ce sont 9 especes qui peuvent étre rapportées
ace genre:
- Blasconura hirtella (Boerner, 1906) sensu Yosii (1976)
= Achorutes hirtellus Boerner, 1906
= Vitronura hirtella; Y osii (1976), Yoshii (1981), Yoshii & Suhardjono (1989)
= Blasconura hirtella, Cassagnau (1983, 1988)
= Vitronura (Blasconura) hirtella; Yoshi & Suhardjono (1992)
espèce décrite de Java et citée d’autres îles indonésiennes (Halmahera et Kei, Yoshii &
Suhardjono, 1992), de Thailande et de Singapour (Yosu, 1976), et de Malaisie (Sabah,
Yoshii, 1981). Les citations du Vietnam et des Philippines reprises par Yoshii &
Suhardjono (1989) d’apres d’anciennes descriptions de Denis (1948) et Gapud (1968)
sont à vérifier (cf. ante). La citation de Taiwan (Yoshii, 1981) demande également
confirmation.
- B. separata (Denis, 1934) comb. n. (Vietnam)
= Achorutes separatus Denis, 1934
- B. anamalensis Cassagnau, 1988 (Inde)
- B. ceylonica Cassagnau, 1988 (Sri Lanka)
- B. palniensis Cassagnau, 1988 (Inde)
- B. prabhooi Cassagnau, 1988 (Inde)
- B. sholica Cassagnau, 1988 (Inde)
- B. toda Cassagnau, 1988 (Inde)
- B. batai sp. n. (Vietnam).
Clé des especes du genre Blasconura
| Tubercules antennaires soudés entre eux et soudés au frontal en une seule
MASSE e Up D)
- Tubercules antennaires soudés entre eux mais séparés du frontal; griffe
Sansidenty st). dele seh gun. BADER ARE LM er ERBOSO O SE 7
2 Soie O absente sur le tubercule antenno-frontal; griffe avec une dent........ 3
- Sole ©) présente sur leitubereule'antenno-frontal «teens 5
3 Ant. I avec 7 soies; macrochètes écailleux; formule des Di: 122/22222
PE a EN CAL HE CL TESTE batai sp. n.
Ant.l'avec'9soles.. 02:45 48e 0 ME NN EEE 4
+ Macrochètes plumeux formule des Diz 133/22223) eee hirtella
- Macrochètes longs et lisses; formule des D:244/53553 eee toda
NOUVEAU COLLEMBOLE NEANURINAE DU VIETNAM 353
5 Griffe sans dent; ant. I avec 8 soies; macrochètes raides, légèrement
HUCUCUK: TOFMULE GES D1:4193/22222 Dir. n ceylonica
- Griffe avec une forte dent; ant. I avec 7 soies; formule des Di: 134/44433 . . . . 6
6 Macrochetes longs, fortement barbelés dans le tiers distal ............: sholica
- Macrochetes’eountsstaiblementecalleur un... me ee prabhooi
7 Soie O absente sur la tête; ant. I avec 7 soies; macrochetes faiblement
nuoueux-ormule des D0122/2292 RR E ee anamalensis
- SOI OMPLESC MCS ET N er A celine OSARE 8
8 Soie O sur les tubercules antennaires soudés; un tubercule supplémen-
taire entre Di et De sur th. II a abd. III; macrochetes fortement barbelés et
renflés à l’apex; formule des Di: 233/33333; ant. I avec 9 soies ..... palniensis
- Soie O sur le tubercule frontal; pas de tubercule supplémentaire sur th. II
a abd. HI; macrochetes ciliés; formule des Di: 333/22222; nombre de
solesssurantl-ineonnu es ae os eee ns Pepe Re ae vee separata
Blasconura batai sp. n.
MATERIEL TYPE. Holotype femelle et 4 paratypes (2 femelles, 1 mâle subadulte et 1
juvénile) en préparation microscopique, | paratype juvénile en alcool. Vietnam: province de Kien
Giang, Hon Chong, colline calcaire de Ba Tai, Berlese de litiere dans un maquis, 19/12/1994
(relevé VIET-034). Holotype et 3 paratypes déposés au Laboratoire d’Ecologie Terrestre
(Université Paul Sabatier, Toulouse, France), 2 paratypes déposés au Muséum d’histoire naturelle
de Geneve (Suisse).
Les abréviations utilisées dans la description sont celles définies par Deharveng
(1983) et Deharveng & Weiner (1984). Le matériel étudié a été récolté par les auteurs.
DESCRIPTION: Longueur: 0,84 a 1,13 mm. Couleur: rouge sur le vivant, blanche
en alcool, yeux non pigmentés. Habitus assez trapu et convexe (fig. 1). Tubercules
dorsaux bien délimités mais peu convexes; grain tertiaire marqué par des groupes de
grains secondaires non soulignés par de véritables réticulations. Abd. VI bilobé, visible
dorsalement. Soies ordinaires dorsales de 2 types: quelques soies fines, lisses et
pointues (mésochètes ou microchetes); macrochetes plus ou moins longs, courbes,
élargis dans leur partie distale en forme de massue, avec une ciliature courte, serrée et
imbriquée, en forme de pousse d’asperge (fig. 2). Soies S fines, atteignant 1/3 (sur le
thorax) a 1/2 (a l’arrière de l’abdomen) de la longueur du macrochète voisin.
Tête (fig. 1, tab Ia et b). Soies S d’ant. IV de taille moyenne, subégales, relati-
vement fines; vésicule apicale nettement trilobée. Ant. I avec 7 soies. Cône buccal fort;
labre ogival assez large, avec 2/2,2 soies (les soies distales longues, dépassant l’apex du
labre). Maxille styliforme, mandibule fine à 3 dents. 2+2 cornéules non pigmentées, la
cornéule antérieure est extérieure et en avant du tubercule oculaire. Tubercules An et Fr
fusionnés, séparés de Oc et de CL. Tubercules Di et De faiblement différenciés, non
fusionnés. Tubercules DL, L et So soudés. Soies O et E absentes. Microchètes Di2 et
De2 sur la bordure interne du tubercule De.
Thorax et abdomen. Tuberculisation et chétotaxie illustrées sur la figure | et le
tableau Ic. Sur abd. IV, le macrochète Di2 est deux fois plus court que le macrochéte
Dil. Dent interne présente sur la griffe. Soie M absente sur le tibiotarse.
354 ANNE BEDOS & LOUIS DEHARVENG
Fics 1-2. Blasconura batai sp. n. 1: habitus et chétotaxie dorsale (longueur du corps: 0,94 mm);
une soie surnuméraire est présente sur le tubercule De de th. II (côté droit); 2: macrochete Del
d’abd. I (longueur de la soie: 73 um).
NOUVEAU COLLEMBOLE NEANURINAE DU VIETNAM 355
REMARQUE: L’holotype présente une anomalie sur th. II (fig. 1). Les 4 paratypes
examinés sont également asymétriques pour la chetotaxie d’au moins un de leurs
tergites. Une forte proportion d’exemplaires non symetriques n’est pas rare chez
certains Neanurinae de la tribu des Lobellini, mais reste exceptionnelle chez les
Paleonurini.
DERIVATIO NOMINIS: Le nom de l’espece est celui de la localité-type, la colline
de Ba Tai, une des rares collines du karst de Hon Chong qui devrait être épargnée par
les carrieres de calcaire.
TABLEAU Ia - Chétotaxie céphalique dorsale de Blasconura batai sp. n.
Tubercule Nombre Type de Soies
de soies soles
CL + 4 ML F
mé G
Af + 8 ML A,B
mi ou mé C,D
Oc + 3 ML Ocm, Ocp
mi Oca
Di + 1 ML Dil
De + 3 ML Del
mi Di2, De2
DL+L+So + 15 ML 4 soies
Mc 1 soie
mé 7 soies
mi 3 soles
TABLEAU Ib - Chétotaxie céphalique ventrale de B. batai sp. n.
Vi 6
Ve 5-6
Labrum 2129)
Labium 10,0x?
Ant. I-II 7,11
Ant. III 1745S
Ant. IV or+8S+i+?mou
Di De DL L Scx2 Cx Tr Fé Ti
Th. I 1 2 1 - 0 3 6 ? 18
Th. I 2. 3+S 3+S+ms 3 2 7 Ÿ ? 18
Th. II À) 4+S 3+S 3 À) 8 ? 11 17
Abd. I 2 3+S 2 3 TV=4+4
Abd. II 2 3+S 2 3 Ve=3-4 (Vel absent)
Abd. III 2 3+S 2 4 Fu=4,0mi Ve=3-4
Abd. IV 2 2+S 3 6 Ve=7 VL=5
Abd. V 2 ——3+S$— (3)-4 Ag=3-(4)+3-(4) VL avec L
Abd. VE —j7 — Ve=14 An=2-3
356 ANNE BEDOS & LOUIS DEHARVENG
DISCUSSION
Deux especes de Blasconura étaient déja connues du Vietnam:
- Blasconura sp., signalée sous le nom de Achorutes hirtellus Boerner, 1906 du “Centre
Annam”, a Bana près de Tourane (Denis, 1934, 1948); il s’agit probablement d’une
espece inedite qui differe du vrai B. hirtella de Java notamment par la présence de 3+S
soies De (au lieu de 2+S) sur abd. IV.
- B. separata (Denis, 1934), signalée sous le nom de Achorutes separatus Denis, 1934
du “Tonkin”, province de Phu-Ho, pres de Yenbai (Denis, 1934, 1948).
Blasconura batai sp. n. est proche de la première de ces espèces mais possède
une chétotaxie dorsale nettement plus réduite. La nouvelle espece partage avec B.
hirtella et B. toda les caractères suivants: la soudure des tubercules An et Fr, l’absence
de soie O sur la tête et la présence d’une dent à la griffe: B. batai sp. n. se distingue de
ces deux especes par plusieurs caracteres chétotaxiques, en particulier la présence de 7
soies sur ant. I au lieu de 9.
REMERCIEMENTS
La campagne de récoltes 1994 a Hon Chong a été possible grâce a l’aide de nos
collègues du département d’Ecologie de l'Université de Ho Chi Minh-Ville et du
Service Science, Technologie et Environnement de la province de Kien Giang, que
nous tenons à remercier.
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NOUVEAU COLLEMBOLE NEANURINAE DU VIETNAM 357
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REVUE SUISSE DE ZOOLOGIE 107 (2): 359-388; juin 2000
Die Leucospidae (Hymenoptera: Chalcidoidea) der Schweiz,
mit einem Bestimmungsschlüssel und Daten zu den
europäischen Arten
Hannes BAUR! & Felix AMIET?
! Abteilung Wirbellose Tiere, Naturhistorisches Museum,
Bernastrasse 15, CH-3005 Bern, Schweiz.
E-mail: hannes.baur@nmbe.unibe.ch
2 Forststrasse 7, CH-4500 Solothurn, Schweiz.
The Leucospidae (Hymenoptera: Chalcidoidea) of Switzerland, with a
key and data on the European species. - In this paper the taxonomy and
distribution of the Swiss species of Leucospis Fabricius are presented. Four
of the seven European species are recorded for Switzerland: L. biguetina
Jurine, L. dorsigera Fabricius, L. gigas Fabricius, and L. intermedia Illiger.
Except for L. dorsigera the main distribution lies in the Southern parts of
the country. The altitudinal gradient ranges from the colline to the
subalpine belt. Adults appear from May to September. L. biguetina and L.
gigas are considered vulnerable with regard to the Swiss fauna. Almost
exclusively females of L. biguetina and L. gigas are known from the study
area, hence these species are likely to reproduce by thelytoky. A key,
diagnoses, morphometric and distributional data are also given for the other
two European species, L. bifasciata Klug and L. brevicauda Fabricius. The
hosts of all species are critically reviewed and new hosts are recorded for
L. dorsigera and L. gigas.
Key-words: Leucospis - taxonomy - key - distribution - new host records -
Switzerland - Europe.
EINLEITUNG
Die Leucospidae stellen mit einer Grösse von ca. 4-16 mm die auffallendsten
Vertreter der zumeist winzigen Erzwespen (Chalcidoidea) dar. Die Familie ist
weltweit mit ca. 140 Arten in 4 Gattungen verbreitet (Boutek, 1974; Noyes, 1998),
wobei in Europa nur gerade 7 Arten der Gattung Leucospis Fabricius vorkommen.
Durch die verdickten Hinterbeine, den über den Gaster nach vorne gebogenen
Legebohrer und die gelb-schwarze Zeichnung sind sie unverkennbar. Alle euro-
päischen Arten führen eine parasitische Lebensweise, d. h. die Larven entwickeln sich
als Ectoparasitoide an Larvenstadien von Wildbienen, z. B. Anthidium, Megachile
oder Osmia (Apidae: Megachilinae) (Clausen, 1940; Bouéek, 1974).
Erste Hinweise auf die Leucospiden der Schweiz finden sich bereits in Sulzer’s
Geschichte der Insekten (1776), wo unter der treffenden Bezeichnung 'Schenkel-
wespe’ nicht wie angegeben L. dorsigera sondern L. gigas aufgrund von Exemplaren
Manuskript angenommen 25.01.2000
360 HANNES BAUR & FELIX AMIET
TABELLE |
Morphologische Terminologie, sortiert nach Abkürzung bzw. Begriff (vgl. Abb. la-Id, gegen-
überliegende Seite).
Ag Auge Auge Ag
Cly Clypeus Clypeus Cly
Col Procoxa Dorsellum Do
Co2 Mesocoxa Gastraltergum Gt
Co3 Metacoxa Mesepisternum Mep
Do Dorsellum Mesocoxa Co2
Fel Profemur Mesofemur Fe2
Fe2 Mesofemur Mesoscutum Msc
Fe3 Metafemur Mesotarsus Ta2
Gt Gastraltergum Mesotibia TH
Mep Mesepisternum Metacoxa Co3
Mpl Metapleuron Metafemur Fe3
Msc Mesoscutum Metapleuron Mp1
Pe Pedicel Metatarsus Ta3
Ppd Propodeum Metatibia Ti3
Prn Pronotum Pedicel Pe
Sca Scapus Procoxa Col
Scr Scroben Profemur Fel
Sct Scutellum Pronotum Prn
Tal Protarsus Propodeum Ppd
Ta2 Mesotarsus Protarsus Tal
Ta3 Metatarsus Protibia Til
Til Protibia Scapus Sca
Ti2 Mesotibia Scroben Ser
Ti3 Metatibia Scutellum Sct
aus Genf beschrieben und illustriert wird. Fuessly (1783) meldet in seinem Beitrag
zur Lebensweise von ‘L. dorsigera’ ebenfalls Funde von L. gigas aus Genf und der
Gegend von Zürich. Etwas später beschreibt Jurine (1807) in seinen «Nouvelle
méthode de classer les hyménopteres et les dipteres» neu L. biguetina Jurine aus der
Schweiz (vgl. Boutek, 1974), daneben erwähnt er auch die Arten L. dorsigera und L.
gigas. Hagenbach (1822) stellt aus der Umgebung von Basel L. fuesslini Hagenbach
auf, ein Synonym von L. dorsigera (vgl. Bouéek, 1974). Labram & Imhoff (1836)
beschreiben und illustrieren letztere und fügen an, dass in der Schweiz noch 2-3
weitere Arten vorkämen. Von Chevrier (1870) erhalten wir unter L. lepida Chevrier
eine weitere Fundortsangabe von L. dorsigera (vgl. Bouëek, 1974) aus dem Genfer-
seegebiet. Schliesslich zählt Schletterer (1890) in seiner bedeutenden, weltweiten
Revision der Leucospiden eine Reihe neuer Funde für die erwähnten Arten ın der
Schweiz auf und führt zusätzlich L. intermedia für das Wallis an. Spätere Arbeiten (z.
B. Boutek, 1959, 1974; Madl, 1990; Schmiedeknecht, 1930) enthalten kaum neue
Angaben und zitieren zumeist in allgemeiner Form aus älteren Schriften.
Während wir recht gut über die Fauna vieler europäischer Ländern unterrichtet
sind (Boucek, 1959, 1964, 1970, 1977; Berland, 1934b; Madl, 1989, 1990; Pagliano,
1998; Schletterer, 1890; Schmid-Egger, 1995), fehlte es bislang an einer
übersichtlichen Darstellung der Leucospiden der Schweiz. Zudem förderte allein die
grobe Durchsicht der Sammlungen von wenigen Museen ein reiches, unpubliziertes
LEUCOSPIDAE (HYMENOPTERA) DER SCHWEIZ 36]
Kopfbreite
Metafemurlänge
Flagellum
1c
Gaster
Ovipositorscheiden
ABB. la-d, L. gigas © : Kopf frontal (a), Metafemur lateral (b), Antenne lateral (c), Körper ohne
Kopf lateral (d) (vgl. Tab. 1, gegenüberliegende Seite). Massstab | mm.
Material aus dem Gebiet zutage. Mit der vorliegenden Arbeit wird erstmals die
Verbreitung der Arten in der Schweiz umfassend dargestellt. Hierzu wurde das
Material von sämtlichen grösseren Sammlungen der Schweiz und mehreren Museen
im Ausland systematisch aufgearbeitet. Die Daten wurden im Hinblick auf eine
Gefährdung der Arten in der Schweiz ausgewertet. Aufgrund von Funddaten der
Wirte wurde versucht, den Grad der Übereinstimmung in der Verbreitung von Wirt
und Parasitoid in der Schweiz aufzuzeigen. Ferner wurde für alle europäischen Arten
die Literatur nach Wirtsangaben durchgesehen, welche kritisch analysiert und durch
362 HANNES BAUR & FELIX AMIET
neue Daten ergänzt wurden. Schliesslich werden ein Bestimmungsschlüssel,
Diagnosen und morphometrische Daten zu allen europäischen Arten vorgelegt.
MATERIAL UND METHODEN
Die vorliegende Studie wertet Daten von insgesamt 919 Exemplaren aus der
Schweiz (572) und Teilen von Europa, dem Nahen Osten und Nordafrika (347) aus.
Für die Verbreitungskarten und die Diagramme zur Höhenverbreitung sowie Phäno-
logie wurde ausschliesslich Material aus der Schweiz verwendet. Den Diagnosen und
Messungen liegt das bei den einzelnen Arten aufgelistete Material zu Grunde. Die
betreffenden Exemplare wurden mit einer Etikette mit dem Vermerk «Baur» und
einer Seriennummer versehen. Messungen wurden an getrocknetem Material unter
einem Stereomikroskop (Leica MZ12) bei 6 (Körperlänge) bzw. 20facher (alle
übrigen Masse) Vergrösserung mit Hilfe eines geeichten Okular-Mikrometers (12 mm
unterteilt in 120 Einheiten) durchgeführt, anschliessend wurden die Werte in Milli-
meter umgerechnet. Die Körperlänge entspricht der Länge vom Vertex bis zur Spitze
des Gasters. Diese Messungen können systematische Effekte (Fehler) aufweisen,
welche von den unterschiedlichen Konservierungsmethoden der einzelnen Sammler
(z. B. die Streckung bzw. Krümmung von Individuen in einer bestimmten Weise)
herrühren. Die Kopfbreite, die Metafemur- und die Ovipositorscheidelänge sind frei
von systematischen Effekten. Die betreffenden Messstrecken sind aus den Abbil-
dungen la, Ib und Id ersichtlich. Der Beginn der Ovipositorscheide (vgl. Abb. 1d) ist
gekennzeichnet durch eine feine Kerbe auf der dorsalen Seite des Ovipositors. Meta-
femur und Ovipositorscheide der linken Seite wurden vermessen. In der morpho-
logischen Terminologie folgen wir Gibson (1997), die verwendeten Begriffe werden
in den Abb. la-1d erläutert (vgl. Tab. 1). Quantitative Angaben beziehen sich auf die
Gesamtheit des betreffenden Materials und sind entsprechend dem System von Goulet
& Mason (1993: 64) wie folgt definiert: sehr selten <1%, selten 1-10%, gelegentlich
>10-25%, manchmal >25-50%, oft >50-75%, häufig >75-90%, meist >90-99%, nor-
malerweise >99% des vorliegenden Materials.
Eine Beurteilung der Gefährdung der einzelnen Arten in der Schweiz muss bei
Parasitoiden, besonders bei streng oligophagen Arten wie Leucospis, naturgemäss die
Gefährdung der Wirte berücksichtigen. Die Angaben richten sich deshalb weitgehend
nach der Roten Liste der gefährdeten Bienen der Schweiz (Amiet, 1994). Zusätzlich
wurde der Anteil rezenter Daten (Funde nach 1950) gewertet. Der Beginn für rezente
Daten wurde auf einen relativ frühen Zeitpunkt festgelegt, da in den letzten drei
Jahrzehnten die Sammeltätigkeit betreffend Leucospiden stark abnahm und sich mehr
oder weniger auf die Anstrengungen von FA beschränkte! Die normalerweise geringe
Populationsdichte sowie die spärliche Besammlung erschweren generell eine Ein-
schätzung des eigentlichen Bestandes. Wenn jedoch eine Art früher bedeutend
häufiger festgestellt wurde als in jüngster Zeit (nach 1950), so kann ein Bestan-
desrückgang angenommen werden (vgl. L. gigas).
Die Daten für das Verbreitungsbild der Wirte stützen sich auf Erhebungen von
FA in folgenden Sammlungen: ETHZ, FA, NMBA, und NMBE. Die Nomenklatur der
mitteleuropäischen Bienen folgt der Liste von Westrich & Dathe (1997). Alle anderen
Namen werden nach Schwarz er al. (1996) zitiert. Die Namen von Blütenpflanzen
LEUCOSPIDAE (HYMENOPTERA) DER SCHWEIZ 363
richten sich nach der «Flora Helvetica» von Lauber & Wagner (1996).
Für die Auswertung der Messreihen bzw. die Erstellung der Verbreitungs-
karten, Grafiken und Abbildungen fand die folgende Software Verwendung: DMAP,
Version 6.5, 1998 (Verbreitungskarten), Adobe Photoshop, Version 5.0.2, 1998
(Beschriftung und Nachbearbeitung der Abbildungen und einiger Diagramme), SPSS,
Version 9.0, 1998 (Auswertung, Diagramme).
Schliesslich sei für alle weiterführenden Angaben zur Synonymie, Morpho-
logie und Literatur der behandelten Arten auf die Monographie der Leucospiden von
Bouéek (1974) sowie den CD-ROM Katalog von Noyes (1998) verwiesen.
Abkürzungen der Kantone der Schweiz
BE Bern so Solothurn
GE Genf TI Tessın
GR Graubünden VD Waadt
NE Neuenburg VS Wallis
SH Schaffhausen ZH Zürich
Abkürzungen der Sammlungen
AS Alı Salvioni, Privatsammlung, Zürich, Schweiz
ASE Albert Sermet, Privatsammlung, Yverdon, Schweiz
BZOL Biologie-Zentrum des Oberösterreichischen Landesmuseums, Linz,
Österreich
(ES) Christian Schmid-Egger, Privatsammlung, Maulburg, Deutschland
DEI Deutsches Entomologisches Institut, Eberswalde, Deutschland
ES Erwin Steinmann, Privatsammlung, Chur, Schweiz
ETHZ Eidgenössische Technische Hochschule, Entomologische Sammlung,
Zürich, Schweiz
FA Felix Amiet, Privatsammlung, Solothurn, Schweiz
FB Frank Burger, Privatsammlung, Orlamünde, Deutschland
FNSM Forschungsinstitut und Naturmuseum Senckenberg, Frankfurt a. M.,
Deutschland
GC Gilles Carron, Privatsammlung, Neuchätel, Schweiz
GR Gerd Reder, Privatsammlung, Flörsheim-Dalsheim, Deutschland
HB Heinrich Bürgis, Privatsammlung, Worms, Deutschland
HT Hansueli Tinner-Guler, Privatsammlung, Landquart, Schweiz
TW Ingmar Wall, Privatsammlung, Mihlingen, Deutschland
MH Martin Hauser, Privatsammlung, Urbana, Illinois, USA [Material
7 Thy (ESS)
MHNG Muséum d’histoire naturelle, Genève, Schweiz
MNHU Museum für Naturkunde, Humboldt-Universität, Berlin, Deutschland
MSNV Museo Civico di Storia Naturale, Venezia, Italien
MZL Musée Zoologique, Lausanne, Schweiz
NMBA Naturhistorisches Museum, Basel, Schweiz
NMBE Naturhistorisches Museum, Bern, Schweiz
NMSO Naturmuseum, Solothurn, Schweiz
NMW Naturhistorisches Museum, Wien, Österreich
364 HANNES BAUR & FELIX AMIET
SMNS _ Staatliches Museum für Naturkunde, Stuttgart, Deutschland
WA Werner Arens, Privatsammlung, Bayreuth, Deutschland
WL Walter Linsenmaier, Privatsammlung, Ebikon, Schweiz
ZSM Zoologische Staatssammlung, München, Deutschland
Bestimmungsschlüssel für die europäischen Arten von.Leucospis (Massstab der
Abbildungen | mm).!
1)— Metatibia dorsal rotbraun bis schwarz, höchstens in der Mitte mit
feinem gelblichem Längsstreifen; basaler Zahn des Metafemurs unge-
fähr so lang wie folgende Zähne (Abb. 2a). d: Gaster im basalen
Drittel stark verschmälert (Abb. 2b); 2: Ovipositorscheide sehr kurz,
nur 0.66-0.74 mal so lang wie der Metafemur............... L. brevicauda
— Metatibia dorsal gelb; basaler Zahn des Metafemurs kürzer oder länger
als folgende Zähne (Abb. 2a', 2a"). d : Gaster im basalen Drittel breiter
(Abb. 2b'); 2: Ovipositorscheide deutlich länger, 0.90-1.93 mal so lang
wieder Metatemüur. u... ee ee LO RR 2
2)— Basaler Zahn des Metafemurs kürzer als folgende Zähne (Abb. 2a'). ....... 3
— Basaler Zahn des Metafemurs länger als folgende Zähne (Abb. 2a") ....... 4
\
2b N 2b'
' Caleca et al. (1995) geben L. miniata Klug für die kleine Insel Lampedusa (Italien)
an. Die vor allem in Nordafrika und dem Nahen Osten verbreitete Art (BouCek, 1974) wurde
im Schlüssel nicht berücksichtigt. L. miniata ist im Habitus sehr ähnlich der orangeroten Form
von L. gigas, sie kann aber leicht durch das gerundete, ungezähnte Dorsellum von den hier
behandelten Arten unterschieden werden.
LEUCOSPIDAE (HYMENOPTERA) DER SCHWEIZ 365
Clypeus stark vorgezogen (Abb. 3a); Flagellum schlank, Glieder 2-4
beim ® länger als breit (Abb. 3b), beim d quadratisch oder wenig
länger als breit (Abb. 3c). Mesoscutum im Zentrum mit 2 gelben
Flecken (Abb. 3d), manchmal diese sich gegenseitig berührend, beim
Gro hte hei dey eee ENTER END. OR ARE L. gigas
Clypeus mässig vorgezogen (Abb. 3a’); Flagellum gedrungener, Glieder
2-4 beim 2 wenig kürzer als breit oder quadratisch (Abb. 3b'), beim 4
deutlich kürzer als breit (Abb. 3c'). Mesoscutum im Zentrum mit nur |
selben Bleck (Abb Sd) bem oitiehlendee Pan aie ae L. intermedia
4) -
HANNES BAUR & FELIX AMIET
Pronotum mit 2 Querkielen: einer im hinteren Viertel, der andere fast
unmittelbar entlang des Hinterrandes (Abb. 4a). 2: Ovipositor erreicht
nicht das 1. Gastraltergum, dieses ohne Längsfurche und sein gelbes
Querband in der Mitte stark eingeschnürt (Abb. 4b); Längsfurche des 5.
Gastraltergums tief, Ovipositorscheide in Ruhelage teilweise verdeckt
CADDRAC) Entei EEE DEE EE L. biguetina
Pronotum mit 3 Querkielen: ein kurzer ungefähr in der Mitte, gefolgt
von zwei längeren im hinteren Viertel respektive fast unmittelbar ent-
lang des Hinterrandes (Abb. 4a'). 9: Ovipositor erreicht das 1. Gastral-
tergum, dieses mit Langsfurche und sein gelbes Querband in der Mitte
nicht eingeschniirt (Abb. 4b'); Langsfurche des 5. Gastraltergums ober-
flachlich, Ovipositorscheide in Ruhelage immer ganz sichtbar (Abb.
ACER RCE Ss RL ee ee RAI N RR REI 0000000 5
LEUCOSPIDAE (HYMENOPTERA) DER SCHWEIZ 367
5)— Clypeus deutlich vorgezogen, lateral mit aufgebogenen Lappen, welche
nach unten leicht konvergieren (Abb. 5a). 9: Längsfurche des 1.
Gastraltergums an der Basis kaum verjüngt (Abb. 5b)......... L. dorsigera
— Clypeus kaum vorgezogen, mehr oder weniger verrundet und lateral
nur mit fein angedeuteten Kielen (Abb. 5a'). 9: Längsfurche des 1.
Gastraltergums an der Basis deutlich verjüngt (Abb. 5b')....... L. bifasciata
1! [orme ne =]
0 5a
rer]
i
en 5b 5b'
Leucospis gigas Fabricius, 1793 Abb. la-Id, 2a', 3a-3d, 6-8, 12-14. Tabl.
Untersuchtes Material [Schweiz: alphabetisch geordnet nach Kanton, Ort; übrige
Länder: alphabetisch geordnet nach Land, Provinz bzw. Region, Ort]: SCHWEIZ: ohne
Fundortsangabe 2 ? (MNHU); «Cornet»: 1 ©, 1867 (MHNG). BE: Bern: 3 2, leg./coll. M.
Perty (NMBE). GE: Genève: 3 ©, leg. Bües (ZSM); 1 ©, 15.6.1889, 2 2, 20.7.1889, 5 9,
20.7.1889, leg./coll. F. Chevrier (MHNG). Genève, Champel: 1 9, [?18...], 1 do, 7.[?18...], 1
99187.018.),1°2,227.048°2] (MENG). GR? Brusio: 1° 99317.3:.8.1933 Je gIc0ll aA
Nadig (ETHZ). VD: Chamblon, Champ Muraz: 1 2, 1942, ex Megachile parietina, leg. A.
Sermet (ASE). Nyon: 3 ©, leg./coll. F. Chevrier (MHNG). VS: 1 2 (MSNV); 1 ®, coll.
Naumberg (DEI); 1 9, leg./coll. FE. Frey-Gessner (NMSO); 114 ®, leg./coll. M. Paul (NMW);
1 ©, leg./coll. O. Schmiedeknecht (NMBE); 1 ®, leg./coll. H. Tournier (MHNG); 6 2, leg. T.
Steck (NMBE); 1 ®, leg. A. Wullschlegel (NMBE). Brig: 2 ®, 7.1914, leg./coll. A. von
Schulthess (ETHZ). Euseigne: 9 2, 8.7.1908, 2 ©, 5.7.1911, leg. T. Steck (NMBE); 1 ©,
87.1908, 1S 2561924 <1 2.,16:8:1929,172157.1936, 1es/collÜT. Steck (NMBA) 339 27
31.7.1939, 1 ©, 23.7.-2.8.1942, leg./coll. A. Nadig (ETHZ); 1 ?, 2.8.1940, leg./coll. J. de
Beaumont (MZL). Finges: 1 2, 25.7.1946, leg./coll. J. Aubert (MZL); Fully, Les Follatéres: 1
2, 5.7.1947, leg./coll. J. Aubert (MZL). Gampel, Jeizinen: 1 9, 8.8.1998, an Sedum album, leg.
H. Baur (NMBE). Martigny: 1 9, 7.7.1935, leg. P. Bovey (MZL); 1 2, 18.7.1936, 1 9
25.6.1948, 1 2, 4.7.1948, 1 9, 28.6.1956, leg./coll. J. de Beaumont (MZL). Niouc: 1 9,
17.7.1900, leg./coll. E. Frey-Gessner (MHNG). Pfynwald: 1 9, 18.6.1921, 1 2, 2.7.1921, leg.
122 Steck (NMBE)FE 27218:641921721725,3:741922 127261924 Ona 702 SMS
20.8.1925, 1 2, 21.7.1927, 2 2, 23.7.1927, leg./coll. T. Steck (NMBA); 6 9, Ende 7.1939, 1 2,
368 HANNES BAUR & FELIX AMIET
13.8.1941, 9 2, 23.7.-2.8.1942, leg./coll. A. Nadig (ETHZ). Salgesch: 1 ©, 11.7.1885,
leg./coll. M. Paul (NMW). Savièse: 1 2, 7.-8.1906, leg. J. Jullien, coll. V. Delucchi (ETHZ);
10 2, 7.1906, leg./coll. J. Jullien (MHNG). Sierre: 3 9, coll. A. von Schulthess (ETHZ); 1 ©,
leg./coll. Cerutti (MZL); 1 9, leg./coll. O. Schmiedeknecht (DEI); 3 2 (MHNG); 1 ©, 6.7.
(MHNG); 3 ©, 25.6.1880, leg./coll. E. Frey-Gessner (ZSM); 1 9, 9.7.1884, 6 2, 7.1886, leg.
T. Steck (NMBE); 2 2, 10.7.1886, leg./coll. R. Meyer-Dür (NMBE); 2 ©, 26.2.1901, coll. A.
von Schulthess (ETHZ); 2 2, 8.1965, leg. I. Wall (IW [nicht untersucht]). Sion: 1 9, leg./coll.
E. Frey-Gessner (MHNG); 3 ®, 25.6.1901, coll. A. von Schulthess (ETHZ); 1 2, 1906, leg. J.
Jullien (MHNG); 1 9, 13.6.1906, leg. Suereuse (MHNG); 1 ®, 1909, leg./coll. B. Jacob
(MZL); 1 2, 25.6.1912, leg./coll. A. von Schulthess (ETHZ); 3 ©, 20.6.1922, 1 9, 28.6.1924,
leg./coll. T. Steck (NMBA). St. Niklaus: 1 2, 15.7.1925, 2 2, 16.7.1925, 1 2, 19.7.1927,
leg./coll. E. Däniker (NMBE); 1 2, 19.7.1927, leg./coll. E. Däniker (WA). Stalden: 1 9,
26.0.1909, leg: T. Steck (NMBE); 1 2, 11.7.1923,.2 9) 14.7.1923 Ales/colSEFDaniker
(NMBE); 1 2, 4.7.1924, leg./coll. T. Steck (NMBA). Susten: 2 9, 24.7.1886, leg./coll. M. Paul
(NMW); 3 2, 13.7.1899, 1 2, 13.7.1899, leg./coll. B. Jacob (MZL); 1 9, 1.7.1900, leg./coll. E.
Frey-Gessner (MHNG); Törbel: 1 2, 8.7.1924, leg. G. C. Krüger (ETHZ); Vex: 1 2, 5.7.1928,
leg./coll. T. Steck (NMBA); Zeneggen: 1 2, 15.-29.7.1943, leg./coll. J. de Beaumont (MZL).
ZH: Zürich: 1 ©, coll. A. von Schulthess (ETHZ); 1 9, «Chalicodoma muraria Nest», leg./coll.
A. von Schulthess (ETHZ); 1 2, 9.1871, leg. F. Ris, coll. A. von Schulthess (ETHZ).
FRANKREICH: Basses Alpes, Digne 1 © (ZSM); Basses Alpes, Valensole 2 2 (NMBE),
2 2,1 d (FA); Bouches-du-Rhône, Fontvielle, Alpille's 1 © (MH); Provence, Callian 1 9
(NMBA), 2 2 (NMBE); Provence, St-Jean 1 2 (MH). GRIECHENLAND: Hellas, Alt Korinth 1 2
(WA); Hellas, Antikes Samiko 1 2 (WA); Hellas, Epid. Limera 1 2 (WA); Hellas, Kalogria,
Achaia 1 2 (WA); Hellas, Leptokoria 2 2 (FB); Hellas, Mavromati, Ithome 1 & (WA); Hellas,
Olympia, Alfios-Tal 1 9, 1 d (WA); Peloponnes, Neapoli, Kap Malea 1 2 (WA). ITALIEN:
Aostatal, Nus 1 2, 1 4 (ES): Aostatal, Pondel 1 9 (CS): Aostatal, St. Pierre 2 2 (CS); Bozen
2 2 (NMW); Bozen, St. Pauls 1 2 (NMW); Sizilien, Passomartino 1 2 (NMBE); Triest 1 2
(ZSM); 1 2 (NMW); Triest, Conconello (Umg.) 4 2 (NMW); Triest, S. Primus 1 2 (NMW);
Vinschgau, Meran 1 2 (DEI). KROATIEN: Dalmatien, Sibenik (Adriaküste) 6 2, ex Megachile
hungarica (GR); Dalmatien, Sibenik 5 9, ex Megachile parietina (GR); Krk 1 2 (ZSM); Krk 1
36 (ETHZ); Krk, Baska 4 9 (ZSM); Krk, Hvar 1 2 (ZSM); Krk, J. Hvar Jelsa 10 ©, 4 à
(ZSM); Krk, Malinska 5 © (ZSM); Rijeka 2 © (ETHZ). MALTA: 3 2 (NMBE). ÖSTERREICH:
Tirol, Klobenstein 1 2 (NMW); Tirol, «Larche» 1 2 (NMW). TUNESIEN: Sbeitla 1 2 (NMBE).
TÜRKEI: Kayseri, Göreme 1 9 (CS).
Diagnose ®: Körper schwarz mit folgenden Teilen gelb [bei manchen
südeuropäischen und nordafrikanischen Individuen leuchtend orangerot]: Gesicht
beiderseits der Scroben mit je einem Fleck (Abb. 3a); Scapus ausser an Basıs und
Apex, manchmal auch dorsal dunkel: Flagellumglieder 1-3 an der Basis häufig leicht
rötlich; Pronotum vorne mit langem, hinten mit etwas kürzerem Querband, oder das
Sklerit vollständig eingerahmt; Mesoscutum mit 2 Flecken zentral und 2 länglichen
Flecken lateral (Abb. 3d); Scutellum am Hinterrand mit in der Mitte stark einge-
schnürtem Querband; Mesepisternum selten mit einem Fleck; Metapleuron voll-
ständig hell; Metacoxa dorsal mit Basalfleck, ventral gelegentlich mit Apikalfleck;
Pro- und Mesofemur apikal zu 2/3 bis 1/4; Aussenseite des Metafemurs, ausser einem
dunklen Fleck und den Zähnen (Abb. 2a’): Tibien und Tarsen, nur Metatibia ventral
dunkel; Gastralterga 1, 4 und 5 mit breiten Querbändern, welche in der Mitte unter-
brochen sind; Tergum 7 lateral mit länglichen Flecken.
Clypeus stark vorgezogen (Abb. 3a); Flagellum schlank, leicht keulenförmig,
Glieder 2-4 länger als breit (Abb. 3b); Pronotum mit 2 Querkielen: einer im hinteren
Viertel, der andere fast unmittelbar entlang des Hinterrandes (ähnlich L. biguetina,
Abb. 4a): Dorsellum am Hinterrand mässig ausgerandet, die beiden Zähne mehr oder
LEUCOSPIDAE (HYMENOPTERA) DER SCHWEIZ 369
weniger deutlich; basaler Zahn des Metafemurs kürzer als folgende Zähne (Abb. 2a’);
Längsfurche des |. Gastraltergums an der Basis leicht verjüngt; Längsfurche des 5.
Tergums oberflächlich, Ovipositorscheide in Ruhelage immer ganz sichtbar (Abb.
1d); Ovipositorscheide 0.95-1.46 mal so lang wie der Metafemur (Abb. 12), reicht bis
ungefähr zur Hälfte, zum basalen Viertel oder zur Basis des 1. Gastraltergums (vgl.
Variabilität 9).
Körperlänge 7.0-17.2 mm (Abb. 13), Kopfbreite 2.15-3.95 mm (Abb. 14).
Diagnose 4G: Gelbe Zeichnungselemente oft reduziert, die Flecken im
Gesicht und im Zentrum des Mesoscutums oft fehlend; Querbänder auf dem Gaster
durchgehend.
Flagellum etwas gedrungener, Glieder 2-4 quadratisch oder wenig länger als
breit (Abb. 3c). Form des Gasters ähnlich L. dorsigera (Abb. 2b’).
Körperlänge 11.2-14.5 mm (Abb. 13), Kopfbreite 2.85-3.40 mm (Abb. 14).
Variabilität 9: Aufgrund des Verhältnisses Ovipositorscheide- zu
Metafemurlänge (O/M) können drei Formen unterschieden werden (Abb. 12), welche
zusätzlich bis zu einem gewissen Grad in der Zeichnung und der Grösse (Abb. 13, 14)
variieren. Die Formen 'A' und 'B' mit kurzer (O/M = 0.95-1.18, n = 79) respektive
langer (O/M = 1.30-1.46, n = 18) Ovipositorscheide treten häufig zusammmen in
mittel- und südeuropäischen Populationen auf. Im Wallis kommt hingegen nur eine
dritte Form vor, welche in der Länge der Ovipositorscheide (O/M = 1.16-1.30, n =
123) eine intermediäre Stellung einnimmt. Die Unterschiede der Walliser Population
zu den beiden anderen Formen sind hoch signifikant (ANOVA mit Dunnett T3-Test,
F = 992.887, n = 220, p < 0.001). Die Formen 'A' und 'B' wurden schon relativ früh
erkannt und als separate Arten unter eigenen Namen beschrieben (Klug, 1814). Nach
Boucek (1974) sind diese aber eindeutig als Synonyme von L. gigas zu werten.
Verbreitung: Paläarktisch, östlich bis China (Bouéek, 1974); in
Mitteleuropa aus allen Ländern nachgewiesen (vgl. Boucek, 1959; Madl, 1989;
Schletterer, 1890). In der Schweiz wurde die Art ausser im Mittelland bei Bern und
Zürich und am Jurasüdfuss bei Yverdon vor allem im Süden (Genferseebecken, GR,
VS) gefunden (Abb. 6). Nach 1950 wurde L. gigas nur noch sehr selten (0.4% der
Daten, 16% der Fundpunkte auf der Verbeitungskarte) und ausschliesslich im Wallis
gesammelt. Die Verbreitung stimmt recht gut mit derjenigen der potentiellen Wirte
überein, wobei diese in der Schweiz weiter verbreitet sind (Nordschweiz, GE, TI, GR)
(Abb. 6).
Höhenverbreitung: von 375 bis 1505 mii. M. (Abb. 6).
Gefährdung: in der Nordschweiz wahrscheinlich ausgestorben, in der
Südschweiz gefährdet und in der ganzen Schweiz stark gefährdet. Von den poten-
tiellen Wirtsarten wird Megachile parietina (Geoffroy) in der Roten Liste (Amiet,
1994) für die Nordschweiz als vom Aussterben bedroht, für die Südschweiz und für
die ganze Schweiz als gefährdet eingestuft. M. pyrenaica Lepeletier, welche vor allem
in den Alpen, selten im Jura und im Mittelland vorkommt, gilt als nicht gefährdet. Die
Beurteilung der Gefährdung von M. parietina trifft ebenfalls recht gut auf L. gigas zu.
Allerdings ist die Art in der Nordschweiz, wo sie nach Sulzer (1776) und Fuessly
(1783) z. B. im Kanton Zürich mehr oder weniger regelmässig auftrat, wohl überall
ausgestorben. Zudem ist sie im Wallis im Rhonetal stark zurückgegangen. Aufgrund
370 HANNES BAUR & FELIX AMIET
- Höhenverbreitung n=154 Phänologie n=90
2000 4
1500
1000
500
— = : :
0% 25% 50 % 75% Mai Juni Juli August September
ABB. 6, L. gigas: Höhenverbreitung, Phänologie und Verbreitung in der Schweiz; @ Funde nach
1950, O Funde vor 1951, potentielle Wirte (Auswertung von 100 Datensätzen von Megachile
parietina und M. pyrenaica).
der Daten war die Art dort früher wahrscheinlich verbreitet und relativ häufig. Gegen-
wärtig ist sie aber nur noch von einer einzigen Stelle bei Jeizinen bekannt. Vor diesem
Hintergrund muss L. gigas für die gesamte Schweiz als stark gefährdet eingestuft
werden.
Biologie: Zu den Wirten werden die folgenden Angaben gemacht,
welche als gesichert gelten können: Megachile parietina (Amiet, Baur, Forster und
Zettel, pers. Beob. an Nestern im Wallis; Fabre, 1886 sub «Chalicodome des
murailles» oder «Chalicodome des galets»; Bouëek, 1970, Fahringer, 1922, Giraud &
Laboulbène, 1877 und Schletterer, 1890 sub Chalicodoma muraria); M. pyrenaica
(Fabre, 1886 sub «Chalicodome des hangars»; Fahringer, 1922 sub Chalicodoma), M.
sicula (Rossi) (Schletterer, 1890 sub Chalicodoma; Bürgis, 1995). Ferner wurden die
LEUCOSPIDAE (HYMENOPTERA) DER SCHWEIZ 371
ABB. 7: Weibchen von L. gigas schlüpft aus einem Nest von Megachile parietina von Sibenik
(Kroatien). Daneben die Exuvie und die Imago der ebenfalls parasitischen Anthrax ?binotatus
Wiedemann (Diptera: Bombyliidae) (Foto G. Reder).
folgenden Exemplare mit zuverlässigen Wirtsangaben untersucht: FRANKREICH: Kor-
sika, Calvi (Umgebung), Stareso, 24.9.1991, ex M. sicula 1 ®, e. p. 11.-13.6.1993,
1 9, e. p. 1994, leg. H. Bürgis (HB). KROATIEN: Dalmatien, Sibenik (Adriaküste) 6
2, e. p. 14.-22.5.1998, ex M. hungarica (Mocsäry), 5 2, e. p. 22.-29.5.1998, ex M.
parietina, leg. G. Reder (GR). SCHWEIZ: VD, Chamblon, Champ Muraz 1 2, 1942, ex
M. parietina, leg. A. Sermet (ASE). ZH, Zürich 1 9, ex M. parietina, leg./coll. A. von
Schulthess (ETHZ).
Die Nennung weiterer Wirte von L. gigas geht zumeist auf eine einzige, oft
recht alte Quelle zurück. So erwähnt Fahringer (1922) den Schlupf von mehreren
Osmia bicornis (Linnaeus) und einem Exemplar von L. gigas aus einem verlassenen
Nest von M. parietina. Da L. gigas bis zu drei Jahren überliegen kann (Bürgis, 1995),
ist nicht auszuschliessen, dass die Entwicklung zuvor an M. parietina, und nicht an
O. bicornis, stattfand. Mann fand «in Corsica einen tiber nussgrossen, kugeligen Bau,
aus Thon und Sandkörnern hergestellt und an einem Zweige von Cystus salviaefolius
befestigt» woraus Odynerus trifasciatus Fabricius (Vespidae), Osmia ‘coerulea’
[= ?0. caerulescens (Linnaeus)] und L. gigas schlüpften (Schletterer, 1890: 11). Hier
372 HANNES BAUR & FELIX AMIET
ABB. 8: Ausschnitt aus dem Habitat von L. gigas bei Jeizinen im Wallis (Schweiz) (Foto B.
Baur).
Osmia als Wirt anzunehmen, ist zwar naheliegend, aber möglicherweise irreführend.
So passt die Beschreibung des «Bau» vorzüglich auf das Nest von M. sicula, ın
diesem Fall der wahrscheinlichere Wirt. Der Parasitoid könnte M. sicula zu einem
frühen Zeitpunkt attackiert haben, und dann, nach Jahren des überliegens, gleichzeitig
mit der O. caerulescens geschlüpft sein, welche in der Zwischenzeit die verwaisten
Zellen für sich benutzte. Die Wirtsangabe Anthophora garrula (Rossi) (Fahringer,
1922 sub Podalirius) bedarf der Bestätigung, diejenige von Vespula vulgaris (Lin-
naeus) (Rondani, 1873) ist äusserst zweifelhaft.
Brutparasiten (Coelioxys, Stelis) werden gelegentlich als Wirte von Leucospis
erwähnt, konnten aber bisher nicht schlüssig bestätigt werden. Boucek (1974)
berichtet lediglich von einem Exemplar von L. gigas, das in Frankreich zusammen
mit Coelioxys quadridentata (Linnaeus) [= C. conica (Linnaeus)] gezogen wurde.
Amiet (pers. Beob.) beobachtete jedoch im Juli in Sambiase, Kalabrien, Italien, wie
ein Exemplar von L. gigas mehrmals immer wieder an der gleichen Stelle das Nest
einer Megachile sp. (parietina oder sicula) anstach. Die Untersuchung des Nestes
ergab vier Zellen, eine verschimmelte, zwei besetzt mit mehreren Larven von ?Stelis
nasuta (Latreille) und eine mit einer fertig entwickelten Megachile-Larve. Die
Leucospis setzte die Spitze des Ovipositors genau über der letztgenannten Zelle an.
Fabre (1886) schildert in seinem Aufsatz über die Leucospis eingehend das
Verhalten von L. gigas. Eine kurze Zusammenfassung über das Eiablageverhalten
sowie die Ontogenie einschliesslich Abbildungen des Eis und der Larvenstadien
findet sich in Clausen (1940). Bürgis (1995) berichtet, dass L. gigas bis zu mehreren
Jahren im Nest von Megachile sicula ausharren könne. Entgegen der verbreiteten
Annahme (z. B. in Jacobs & Renner, 1998), wonach die fertig entwickelte Wespe auf
LEUCOSPIDAE (HYMENOPTERA) DER SCHWEIZ 373
die Hilfe eines zuvor schlüpfenden Wirtes, welcher einen Ausgang freimache, ange-
wiesen sei, befreiten sich alle Exemplare selbständig aus dem Nest (Bürgis, 1995).
Ferner konnte Reder (pers. Beob.) mehrere Exemplare von L. gigas beim Schlupf aus
Nestern von M. parietina (Abb. 7) bzw. M. hungarica beobachten. Auch in diesem
Fall haben alle Parasitoide mit Hilfe der Mandibeln scheinbar mühelos das Nest
selbständig verlassen.
Imagines erscheinen von Juni bis August (Abb. 6) und werden meist an
xerothermen Standorten wie Steppen- und Trockenrasen (Abb. 8) angetroffen, wo sie
in unruhigem, raschem Flug das Gelände nach Nestern der Wirte absuchen. Einzelne
Exemplare konnten auf Blüten von Allium sphaerocephalon Linnaeus (Liliaceae)
(Forster, pers. Beob.) und Sedum album Linnaeus (Crassulaceae) (vgl. Material)
beobachtet werden. Nach Ansicht verschiedener Autoren (Berland, 1934a; Bytinski-
Salz, 1963) vermehrt sich L. gigas in Europa vor allem parthenogenetisch (thelytok),
da hier Männchen nur selten gefunden werden. Dies scheint auch für die Populationen
in der Schweiz zuzutreffen, denn unter den insgesamt 281 untersuchten Exemplaren
befindet sich nur ein einziges Männchen aus der Nähe von Genf.
Leucospis intermedia Niger, 1807 Abb. 3a'-3d', 9, 12-14
Untersuchtes Material: SCHWEIZ: GR: Brusio: 2 d, 31.7.-5.8.1935, leg./coll. A. Nadig
(ETHZ). Grono: 3 ©, 7.1887, leg. T. Steck (NMBE); 1 2, 21.8.1934 (NMBA). Roveredo: 1 d,
20.7.1928, leg./coll. A. Nadig (ETHZ); 1 2, 7.8.1935, leg./coll. T. Steck (NMBA). VD:
Buchillon: 1 ©, 17.7.1930, leg. P. Bovey (ETHZ). Nyon: 1 6, leg./coll. F. Chevrier (MHNG).
VS: ohne Fundortsangabe 1 9, leg. A. Wullschlegel (NMBE); 1 d, leg./coll. E. Frey-Gessner
(NMW); 1 9, leg./coll. H. Tournier (MHNG). Aven: 1 @, 28.5.1992, leg./coll. F. Amiet
(NMSO). Branson: 2 4, 30.5. (MHNG); 1 d, 23.6.1964, 1 ?, 29.7.1964, leg./coll. J. de
Beaumont (MZL). Euseigne: 1 6, 5.7.1911, leg. T. Steck (NMBE); 2 2, 31.7.1939, leg./coll.
A. Nadig (ETHZ). Evolène: 1 4, 14.7.1911, leg. T. Steck (NMBE). Fully, Les Follatères: 1 4,
1929 (MHNG); 2 2, 11.7.1933, 1 2, 21.7.1933, 1 8,1 2, 27.6.1942, leg. P. Bovey (MZL); 2
221824126194 I 25 195.1948, leg/coll. A. Nadis (EIHZ); 1 2, 14.6.1951, leg./coll. J.
Aubert (MZL); 1 2, 29.7.1964, leg./coll. J. de Beaumont (MZL). Grimentz: 1 ©, 27.7.-
12.8.1944, leg./coll. J. de Beaumont (MZL). Val d'Hérens: 1 4, 26.6.1894, leg./coll. M. Paul
(NMW). Martigny: 1 9, leg./coll. B. Jacob (MZL); 1 3, 1 2, 1.-4.6.1875, 1 9, 7.-8.6.1875,
leg./coll. E. Frey-Gessner (ZSM); 1 8,5 9, 15.6.1889, leg. T. Steck (NMBE); 1 9, 5.7.1933,
leg./coll. J. de Beaumont (MZL). Mazembroz: 1 6, 14.5.1934, leg. P. Bovey (ETHZ); 1 d,
18:541973712.870:0:1976,, 1.8, 15:8:1977, 273, 2.619887 leg. E. Amict (EA) Nioucs 279,
12.8.1908, leg. T. Steck (NMBE). Ried bei Morel, Riederalp: 1 2, 16.7.1921, leg./coll. E.
Däniker (NMBE). Sierre: 1 d, 3 ©, coll. A. von Schulthess (ETHZ); 1 2, leg./coll. A. von
Schulthess (ETHZ); 1 d, leg./coll. O. Schmiedeknecht (NMW); 1 à, leg./coll. O. Schmiede-
knecht (MNHU); 1 4, 16.7.1880, ex Megachile sp., leg./coll. E. Frey-Gessner (MHNG); 1 4,
9.7.1884, leg./coll. A. von Schulthess (ETHZ); 1 d, 22.7.1884, 1 9, 7.1886, leg. T. Steck
(NMBE); 1 d, 24.6.1885, leg./coll. M. Paul (MHNG); 1 d, 27.6.1886, 3 2, 1887, leg. M. Paul,
coll. A. von Schulthess (ETHZ). Sion: 1 2, leg./coll. F. Chevrier (MHNG); 1 6, 8.7.1884, leg.
T. Steck (NMBE); 1 2, 9.1912 (NMBA). St. Niklaus: 1 2, 19.7.1927, leg./coll. E. Däniker
(NMBE). Stalden: 1 6, 26.6.1909, 1 d, 1 2, 27.6.1909, 3 d,2 9, 28.6.1909, leg. T. Steck
(NMBE). Vex: 2 d, 21.6.1935, leg./coll. T. Steck (NMBA). Zeneggen: 1 ©, 20.7.1986, leg. F.
Amiet (FA).
FRANKREICH: Py-Montet 1 d (MH); Tence 1 2, ex Osmia emarginata (MHNG). GRIE-
CHENLAND: Hellas, Kallogria, Achaia 2 £ (WA); Peloponnes, Blafonisoa 1 2 (WA). ITALIEN:
Aostatal, W Arvier 1 & (CS); Aostatal, Sarre 1 £ (CS); Aostatal, St. Pierre 1 2 (CS); Bozen 2
2,13 (NMW); Bozen 2 9, 2 gd (ETHZ); Puglia, Mte. Gargano, NW M. S. Angelo 1 2
(ETHZ); Sizilien, Siracusa 1 2 (NMBE); Vinschgau, Latsch 1 2 (ES). KROATIEN: Dalmatien,
374 HANNES BAUR & FELIX AMIET
Split 6 2 (NMW); Krk 2 ? (ETHZ); Krk, Hvar 2 ® (ZSM); Krk, J. Hvar Jelsa 1 2 (ZSM).
ÖSTERREICH: Kärnten, Villach 1 2 (NMW); Tirol, «Larche» 1 2 (NMW).
Diagnose ® ähnlich der von L. gigas, aber: Flagellumglieder 1-3 nie
rötlich; Pronotum vorne mit kurzem, hinten mit langem Querband (ähnlich L.
dorsigera, Abb. 4a'), oft das Sklerit vollständig eingerahmt; Mesoscutum mit nur 1
Fleck zentral; Propodeum selten mit je einem kleinen Fleck lateral; Querband auf
Gastraltergum 4 manchmal kaum unterbrochen.
Clypeus mässig vorgezogen (Abb. 3a’); Flagellum gedrungener als bei L.
gigas, fast zylindrisch, Glieder 2-4 wenig kürzer als breit oder quadratisch (Abb. 3b’);
Längsfurche des |. Gastraltergums normalerweise nicht verjüngt; Ovipositorscheide
länger, 1.46-1.93 mal so lang wie der Metafemur (Abb. 12), reicht meist mindestens
bis zum Hinterrand des Scutellums, oft bis zu dessen Mitte.
Körperlänge 4.5-12.3 mm (Abb. 13), Kopfbreite 1.45-3.10 mm (Abb. 14).
Diagnose d: Gelbe Zeichnungselemente oft reduziert, der Fleck im
Zentrum des Mesoscutums oft fehlend; Querbänder auf dem Gaster durchgehend.
Flagellum etwas gedrungener, Glieder 2-4 deutlich kürzer als breit (Abb. 3c’).
Form des Gasters ähnlich L. dorsigera (Abb. 2b’).
Körperlänge 6.8-10.2 mm (Abb. 13), Kopfbreite 2.05-2.85 mm (Abb. 14).
Verbreitung: Paläarktisch, östlich bis Tadschikistan (Bouéek, 1974); in
Mitteleuropa aus allen Ländern nachgewiesen (vgl. Boutek, 1959; Madl, 1989, 1990;
Schletterer, 1890). In der Schweiz ist die Art auf das Wallis, das Genferseegebiet und
die Alpensüdtäler beschränkt (Abb. 9). Nach 1950 wurde L. intermedia nur noch
gelegentlich (19% der Daten, 22% der Fundpunkte auf der Verbreitungskarte) und
ausschliesslich im Wallis gesammelt. Die Verbreitung stimmt, mit Ausnahme des
Genferseebeckens, recht gut mit derjenigen des potentiellen Wirtes überein, wobei
dieser in der Schweiz weiter verbreitet ist (Mittelland, Alpen) (Abb. 9).
Höhenverbreitung: von 290 bis 1925 mii. M. (Abb. 9).
Gefährdung: nicht gefährdet. Der potentielle Wirt Osmia mustelina
Gerstäcker hat keinen Eingang in die Rote Liste (Amiet, 1994) gefunden und gilt in
der Schweiz als nicht gefährdet.
Biologie: Als Wirt wird von verschiedenen Autoren bisher nur Osmia
emarginata Lepeletier angegeben (z. B. Berland, 1934b für Frankreich; Bouéek, 1959
für Tschechien; Giraud, 1858 für Österreich; Madl, 1990 für Deutschland). Allerdings
werden in jüngerer Zeit die Populationen östlich von Frankreich als eigene Art, O.
mustelina, aufgefasst. Daher kommt als Wirt von L. intermedia in der Schweiz vor
allem diese Osmia-Art in Frage. Ferner wurden die folgenden Exemplare mit Wirts-
angabe untersucht: FRANKREICH: Haute-Loire, Tence 1 9, 16.7.1925, «parasite d’O.
emarginata» (MHNG); SCHWEIZ: VS, Sierre 1 d, 16.7.1880, «nid de Megachile No.
9320, e. p. 4. 1881», leg./coll. E. Frey-Gessner (MHNG); SPANIEN: Soria, Beraton
1 9, 22.6.1994, an Nest von O. emarginata, leg. F. Amiet (FA). Die Wirtsangabe von
Frey-Gessner bedarf der Bestätigung!
Imagines erscheinen von Mai bis August (Abb. 9) und werden meist an xero-
thermen Standorten wie Steppen- und Trockenrasen angetroffen, wo sie in raschem
Flug das Gebiet nach Nestern ihrer Wirte absuchen. Frühere Vermutungen, dass sich
L. intermedia in Europa vor allem parthenogenetisch (thelytok) fortpflanze (Boucek,
1964), konnten nicht bestätigt werden, da Männchen überall recht zahlreich gefunden
wurden.
LEUCOSPIDAE (HYMENOPTERA) DER SCHWEIZ 375
Höhenverbreitung n=84 50%, Phanologie n=62
25% +
0%
r 7
50 % 75% Mai Juni Juli August September
SEI je à
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2) à
ABB. 9, L. intermedia: Höhenverbreitung, Phänologie und Verbreitung in der Schweiz; @ Funde
nach 1950, © Funde vor 1951, potentieller Wirt (Auswertung von 38 Datensätzen von Osmia
mustelina).
Leucospis dorsigera Fabricius, 1775 Abb. 2b’, 4a'-4c', Sa, Sb, 10, 12-14
Untersuchtes Material: Schweiz: ohne Fundortsangabe 1 4, leg. Morawitz, coll. Gerst.
(MNHU). BE: Batterkinden: 1 2, 25.7.1887, leg./coll. T. Steck (NMBA). Bern: 2 2, leg./coll.
M. Perty (NMBE); 1 9, 1.8.1897, leg. T. Steck (NMBE). Bern, Kirchenfeld: 1 3, 9.6.1922,
leg./coll. T. Steck (NMBA). Biel: 1 2, 29.7.1888, leg./coll. T. Steck (NMBA). Lyss: 1 2,
26.6.1908, leg. T. Steck (NMBE); 1 ®, 26.8.1923, leg./coll. T. Steck (NMBA). La Neuveville:
1 2, 13.7.1897, leg./coll. B. Jacob (MZL). GE: l'Allondon: 1 2, 29.6.1957, 2 2, 20.6.1959,
leg./coll. Steffen (MHNG). Genève, Champel: 1 2, 17.9.[?18...] (MHNG). Genève, Corsier: 1
2, 14.7.1985, leg./coll. Steffen (MHNG). Russin: 1 ©, 6.8.1995, leg. F. Amiet (FA). Satigny,
Peney: 1 9, 30.5.1875 (MHNG); 1 6, 21.6.1875 (MHNG); 1 ©, 7.1875 (MHNG); 1 28,
1.7.1875 (MHNG). GR: Grono: 4 2, 7.1885, leg. T. Steck (NMBE); 1 9, 27.6.1921, 2 ®,
30.6.1923, 1 2, 19.8.1924, leg./coll. T. Steck (NMBA). Klosters: 1 £ (ETHZ). Landquart: 1
3, 8.6.1996, 1 2, 14.6.1996, an Osmia adunca Brutplatz, leg. H. Tinner-Guler (HT). Mesocco:
1 2, 14.7.1927, leg./coll. T. Steck (NMBA). Roveredo: 1 2, 19.8.1924, 1 9, 18.-20.6.1926,
376 HANNES BAUR & FELIX AMIET
1 2720741928,1 8,2 2, 23.7.-3.8.1933, 1 2, 17.8.1949 IegJeoll. Ay Nadig (ERHZ) MINCE
1 2, 18.8.1934, 1 gd, 5.8.1935, 1 2, 7.8.1935, leg./coll. T. Steck (NMBA). Roveredo, S.
Vittore: 1 2, 3.-10.8.1942, leg./coll. A. Nadig (ETHZ). NE: Auvernier: 1 ©, 7.9.1907, leg./
coll. B. Jacob (MZL). SH: Merishausen: 1 9, 9.8.1999, leg./coll. G. Bächli (NMBC). SO:
Dornach: 1 9, 3.8.1995, leg./coll. F. Amiet (FA). Solothurn: 1 ©, 4.8.1995, leg./coll. F. Amiet
(FA). TI: Acquarossa: 1 2, 9.-10.1951, leg./coll. Lautner (NMBA). Agno: 2 6, 1 9,
15.7.1944, leg./coll. J. de Beaumont (MZL). Bignasco: 1 2, 1905, leg. T. Steck (NMBE).
Capolago: 1 9, 12.7.1944, leg./coll. J. de Beaumont (MZL). Chiasso, Pedrinate: 1 9,
16.8.1997, leg. F. Amiet (FA); Claro: 1 ®, 6.8.1996, leg. A. Salvioni (AS). Gordola: 1 à,
26.7.1933, leg./coll. A. Nadig (ETHZ). Losone: 1 2, 30.6.-2.7.1959, leg./coll. J. de Beaumont
(MZL). Maroggia: 1 6, 8.[719...], 1 2, 27.8.[719...], 1 2, 8.1919, leg. G. C. Krüger, coll. A.
von Schulthess (ETHZ). Minusio: 1 d, 4 ®, 30.6.1953, 1 ©, 30.6.1953, an Apiaceae, 2 9,
7.7.1953, leg./coll. A. Nadig (ETHZ). Novaggio: 1 2, 6.1911, coll. A. von Schulthess (ETHZ);
1 d, 26.6.1920, leg./coll. T. Steck (NMBA). VD: Bonvillars, 1 9, 3.8.1981, leg./coll. Steffen
(MHNG). Commugny: 1 ®, 24.7.1949, 1 2, 1.8.1954, 1 2, 21.7.1948, 1 2, 6.8.1950, 1 2,
19.7.1949, 1 9, «Eté» 1946, 1 ?, 24.7.1955, leg./coll. Steffen (MHNG). Cudrefin: 1 9,
15.8.1926, leg./coll. T. Steck (NMBA). La Sauge: 1 ?, 10.8.1959, leg./coll. J. de Beaumont
(MZL). VS: 1 à, coll. A. von Schulthess (ETHZ): 1 d, coll. H. Tournier (MHNG). Chandolin:
1 2, 5.7.1968 (MHNG); 1 9, 7.7.1968 (MHNG). Euseigne: 1 2, 3.7.1897, 1 2, 6.8.1933, leg.
T. Steck (NMBE); 1 9, 27.6.1924, 1 9, 6.8.1932, 1 2, 8.8.1932, leg./coll. T. Steck (NMBA);
1 3, 25.-26.7.1939, 1 gd, 12.6.1948, leg./coll. A. Nadig (ETHZ). Fully, Les Follatéres: 1 à,
23.7.-2.8.1942, leg./coll. A. Nadig (ETHZ). Hohtenn: 1 ®, 29.9.1987, 1 3, 15.9.1991, leg. F.
Amiet (FA); 1 ©, 26.7.1994, leg. S. Ungricht (ETHZ). Leuk, Platten, Rotten: 1 2, 10.8.1997,
leg. B. Merz (ETHZ). Martigny: 1 6, 15.6.1889, 1 d, 15.6.1889, leg. T. Steck (NMBE); 1 6,
18.7.1936, 1 ©, 29.6.1946, leg./coll. J. de Beaumont (MZL). Mazembroz: 1 ®, 6.6.1976, leg.
F. Amiet (FA). Morel: 1 ©, 1.8.1939, leg./coll. A. Nadig (ETHZ). Niouc: 1 d, 12.7.1908, leg.
T. Steck (NMBE). Pfynwald: 1 d, 28.8.1885, leg. M. Paul, coll. A. von Schulthess (ETHZ); 1
2, 9.7.1975, leg. F. Amiet (FA); 1 2, 15.7.1977, leg. F. Amiet (FA); 1 ©, 1989, leg. M. Paul,
coll. A. von Schulthess (ETHZ); 3 2, 13.8.1997, leg. B. Merz (ETHZ). Sierre: 2 9, coll. A.
von Schulthess (ETHZ); 1 à, 7.7.[718...], 1 ©, 1.8.[218...] (MHNG); 1 ©, 20.7.1884, les. T.
Steck (NMBE); 3 ®, 1887, 2 9, 7.9.1887, leg. M. Paul, coll. A. von Schulthess (ETHZ).
Sierre, Le Glarey: 1 9, 12.7.1905, leg. J. Jullien (MHNG). Sion: 1 2, 25.6.1912, leg./coll. A.
von Schulthess (ETHZ). St. Niklaus: 1 ©, 15.7.1925, leg./coll. E. Däniker (NMBE). Susten: 1
2, leg./coll. B. Jacob (MZL); 1 ®, 7.8.1887, leg. M. Paul, coll. A. von Schulthess (ETHZ). ZH:
Zürich; 1 2, coll. A. von Schulthess (ETHZ); 1 2, 9.1870, leg. F. Ris, coll. A. von Schulthess
(ETHZ). Zürich, Albisgiietli: 1 9, 31.7.1995, an Daucus carota, leg. S. Ungricht (ETHZ).
DEUTSCHLAND: Baden-Wiirttemberg, Freiburg i. B. 1 2, 2 d (ETHZ); Baden-Wiirttem-
berg, Kaiserstuhl 4 2, 1 & (ETHZ); Bayern, Dudenhofen 1 & (MH); Bayern, Eichstätt 2 9
(ZSM); Bayern, Nürnberg, Laufamholz 1 2,2 4 (ZSM); Hessen, Flörsheim-Dalsheim 1 9
(GR); Landau 1 © (ZSM); Thüringen, Frankenhausen, Kyffhäusern 2 © (DEI). FRANKREICH:
Basses Alpes, Valensole 1 £ (NMBE); Bouches-du-Rhône, St-Martin de Crau 1 2, 1 4 (MH);
Gard, Nimes 1 2 (ZSM); Provence, Callian 1 2 (NMBE); Roussillon, Argelès-sur-Mer 1 2,
1 3 (FA). ITALIEN: Abruzzi, L'Aquita, Fonte Cerreto 1 d (ZSM); Aostatal, Sarre 1 9 (CS);
Bozen 1 d (NMBA), 1 2 (DEI), 10 2,8 d (NMW); Bozen, St. Pauls 23 2 (NMW); Lavagna
1 2 (ZSM); Lecco, Mte. Baro 1 9 (ETHZ); Lecco, Pusiano 12 2 (ETHZ); Sizilien, Siracusa
1 2 (NMBE); Triest, Conconello 2 2 (NMW); Vinschgau, Meran 1 9 (ZSM). KROATIEN: Krk,
Baska 2 2, 2 6 (ZSM); Krk, Hvar 3 2, 3 6 (ZSM); Krk, Rudina 2 2 (ZSM). ÖSTERREICH:
Burgenland, Winden 29 9, 5 4 (NMW); Kärnten, Podcetrtek 1 2 (ZSM); Niederösterreich,
Furth 1 ¢ (NMW); Oberösterreich, Linz 1 d (NMW); Salzburg, Glau-Moos am Heustadl 1 9,
3 d (ZSM); Salzburg, Porsch 1 2 (ZSM); Wien, Mauer 2 & (NMW). SLOWENIEN: Ljubljana
[Laibach, Carniolia] 3 2,1 & (ZSM).
Diagnose 9: Körper schwarz mit folgenden Teilen gelb: Scapus von
dunkel bis ganz hell; Pronotum vorne mit kurzem, hinten mit langem Querband (Abb.
4a‘), manchmal das Sklerit vollständig eingerahmt; Mesoscutum selten mit 2 läng-
LEUCOSPIDAE (HYMENOPTERA) DER SCHWEIZ 377
lichen Flecken lateral; Scutellum am Hinterrand mit schmalem bis mässig breitem
Querband; Metapleuron teilweise bis vollständig hell; Pro- und Mesocoxa vereinzelt
mit kleinem Fleck, Metacoxa ventral oft mit variablem Apikalfleck; Pro- und
Mesofemur apikal zu 1/3 bis 1/10, Metafemur meistens basal und apikal mit je einem
Fleck, diese manchmal breit verbunden; Tibien und Tarsen grösstenteils: Protibia oft
lateral, Metatibia immer ventral dunkel; Gastralterga 1, 4, 5 und 7 mit variabler
Zeichnung (Abb. 4c'): Terga 1 und 5 mit breitem Querband, beim ersten durch die
Längsfurche unterbrochen (Abb. 5b); Terga 4 und 7 mit lateralen Flecken, welche
manchmal reduziert sind und auf Tergum 4 gelegentlich sogar fehlen.
Clypeus deutlich vorgezogen, lateral mit aufgebogenen Lappen, welche nach
unten leicht konvergieren (Abb. 5a); Flagellum schlank, leicht keulenförmig, Glieder
2-4 quadratisch oder länger als breit; Pronotum mit 3 Querkielen: ein kurzer ungefähr
in der Mitte, gefolgt von zwei längeren im hinteren Viertel respektive fast unmittelbar
entlang des Hinterrandes (Abb. 4a’); Dorsellum am Hinterrand nur leicht ausgerandet,
Zähne daher undeutlich; basaler Zahn des Metafemurs breit, deutlich länger als die
folgende Reihe sehr kleiner Zähne (ähnlich L. biguetina, Abb. 2a"); Längsfurche des
1. Gastraltergums an der Basis kaum verjüngt (Abb. 5b); Längsfurche des 5.
Gastraltergums oberflächlich, Ovipositorscheide in Ruhelage immer ganz sichtbar
(Abb. 4c'); Ovipositorscheide 1.53-1.92 mal so lang wie der Metafemur (Abb. 12),
reicht normalerweise mindestens bis zur Basis des |. Tergums, oft bis zum Scutellum.
Körperlänge 5.7-13.2 mm (Abb. 13), Kopfbreite 1.60-3.15 mm (Abb. 14).
Diagnose d: Gelbe Zeichnungselemente mässig bis stark reduziert,
kleine Exemplare gelegentlich bis auf helle Stellen an Pronotum, Beinen und Gaster
vollständig dunkel; Querbänder auf dem Gaster oft durchgehend.
Flagellum leicht gedrungener. Gaster im basalen Drittel leicht verschmälert
(Abb. 2b’).
Körperlänge 5.0-10.5 mm (Abb. 13), Kopfbreite 1.35-2.55 mm (Abb. 14).
Verbreitung: Paläarktisch, östlich bis Tadschikistan (Bouëek, 1974); in
Mitteleuropa aus allen Ländern nachgewiesen (vgl. Boucek, 1959; Madl, 1989;
Schletterer, 1890; Schmid-Egger, 1995; Schmidt, 1969; Wolf, 1953). In der Schweiz
kommt die Art an wenigen Stellen im Jura, Mittelland und Graubünden und an recht
zahlreich Orten in der Südschweiz (GE, VS und TI) vor (Abb. 10). Nach 1950 wurde
L. dorsigera nur manchmal (26% der Daten, 42% der Fundpunkte auf der Ver-
breitungskarte) gesammelt. Die Verbreitung stimmt recht gut mit derjenigen der
potentiellen Wirte überein, wobei diese zusammen ein deutlich grösseres Gebiet
besiedeln (z. B. die Alpen und das Engadin) (Abb. 10).
Höhenverbreitung: von 240 bis 1125 m i. M. (Abb. 10).
Gefährdung: nicht gefährdet. Die meisten der potentiellen Wirte (vel.
Biologie) werden in der Roten Liste (Amiet, 1994) als nicht gefährdet aufgeführt.
Biologie: Aus Nestern von Anthidium diadema Latreille (Fabre, 1886), A.
strigatum (Panzer) (Saunders, 1875 sub A. contractum), Osmia adunca (Panzer)
(Amiet, pers. Beob.; Grandi, 1961; Müller er al., 1997; Westrich, 1989), O. bicornis
(Linnaeus) (Boucek, 1970; Le Goff, 1999 und Schletterer, 1890 sub O. rufa), O.
fedtschenkoi (Morawitz) (Herting, 1977), O. niveata (Fabricius) (Herting, 1977 und
Wolf, 1953 sub O. fulviventris) und O. tricornis Fabricius (Le Goff, 1999) gezogen.
Ferner wurde das folgende Exemplar mit zuverlässigen Wirtsangaben untersucht:
378 HANNES BAUR & FELIX AMIET
Höhenverbreitung n=122 50% = Phänologie n=93
2000 |
1500
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ABB. 10, L. dorsigera: Höhenverbreitung, Phänologie und Verbreitung in der Schweiz;
@ Funde nach 1950, O Funde vor 1951, potentielle Wirte (Auswertung von 213 Datensätzen
von Anthidium strigatum, Osmia adunca, O. bicornis, O. ligurica und O. niveata). Der Fundort
bei Merishausen (SH) konnte auf der Verbreitungskarte nicht mehr berücksichtigt werden.
ITALIEN: Toskana, Volterra 1 9, 10.1985, ex Nest Osmia ligurica Morawitz, leg. F.
Amiet (FA).
Reder (pers. Beob.) fing in seinem Garten ein Weibchen an Bruthölzern, worin
seit vielen Jahren Osmia cornuta (Latreille), O. parietina Curtis, O. bicornis und
Megachile ericetorum Lepeletier brüteten. Grandi (1961) beobachtete L. dorsigera an
alten Gängen von Bostrichiden und Cerambyciden (Coleoptera), die von Bienen der
Gattung Eriades [= Osmia] besiedelt waren. Die Angabe von Bostrichus monacha
(Fabricius) (Coleoptera: Bostrichidae) in Herting (1973) ist gewiss ebenfalls auf die in
den Gängen des Käfers nistende Biene zu beziehen.
Imagines erscheinen von Mai bis September (Abb. 10) und werden meist an
offenen, sonnigen Standorten wie Trocken- und Halbtrockenrasen, aber auch grösse-
ren Waldlichtungen angetroffen. Einzelne Exemplare konnten auf Blüten von Daucus
LEUCOSPIDAE (HYMENOPTERA) DER SCHWEIZ 379
carota Linnaeus und anderen Doldengewächsen (Apiaceae) sowie Euphorbia (Eu-
phorbiaceae) beobachtet werden (vgl. Material; Amiet, Burger, Merz, pers. Beob.;
Chevrier, 1870; Pagliano, 1998). Aufgrund der zahlreich nachgewiesenen Männchen
darf angenommen werden, dass sich L. dorsigera in der Schweiz in erster Linie
bisexuell (arrhenotok) fortpflanzt.
Leucospis biguetina Jurine, 1807 2a", 4a-4c, 11-14
Untersuchtes Material: Lectotypus 9, ohne Fundortsangabe [Herkunft nach Boucek
(1974): Schweiz], coll. L. Jurine (MHNG). Schweiz: VS: ohne Fundortsangabe: I 2, coll.
Naumberg (DEI). Ausserberg: 1 £, 29.7.1933, leg. M. Naef (WL). Chandolin, Soussillon: 2 9,
30.6.1921, leg./coll. T. Steck (NMBA). Euseigne: 1 9, 14.8.1933, leg. M. Naef (WL).
Grengiols: 1 ©, 22.6.1998, leg./coll. F. Amiet (NMBE). Grimentz: 1 ©, 17.7.1978, leg./coll. F.
Amiet (NMBE). Hérémence: 1 9, 23.7.1992, leg. F. Amiet (FA). Hohtenn: 2 2, 10.8.1930, 2
2, 21.8.1932, 1 2, 29.7.1934, 2 2, 7.7.1935, leg. M. Naef (WL); 1 2, 29.7.1987, leg. F. Amiet
(FA); 1 ®, 29.7.1987, leg./coll. F. Amiet (NMBE). Hohtenn, Ladu: 1 ©, 15.7.1991, leg. F.
Amiet (FA). Lens: 1 9, 11.7.1967, 1 ®, 13.7.1967, 4 2, 16.7.1967, an Apiaceae (MHNG); 1
2, 22.7.1927, leg. T. Steck (NMBE). Martisberg: 1 2, 21.7.1992, leg./coll. F. Amiet (NMBE).
Niedergesteln: 1 9, 8.8.1998, leg. F. Amiet (FA). Niouc: 2 9, 12.7.1908, leg. T. Steck
(NMBE). Pfynwald: 1 9, 29.6.1924, leg./coll. T. Steck (NMBA); 1 @, 18.7.1958, 1 2,
2.8.1960, leg. Daicker (MHNG); 1 2, 7.1958, 1 ©, 10.7.1975, leg./coll. F. Amiet (NMSO); 1
M0 7107502 77-1975 1,2 ,8:7.1979 leg; F'Amet (FA) M MO T7 197ÉMeL colle:
Amiet (NMBE); 1 @, 6.7.1997, leg. B. Merz (NMBE); 1 2, 12.8.1997, leg. B. Merz (ETHZ).
Sierre: 2 ©, coll. A. von Schulthess (ETHZ); 1 ©, leg./coll. O. Schmiedeknecht (DEI); 2 9,
leg. O. Schmiedeknecht (NMW); 1 9, leg./coll. O. Schmiedeknecht (MNHU); 1 2, leg./coll.
O. Schmiedeknecht (ETHZ); 1 2, leg./coll. T. Steck (NMBA); 1 ® (DEI); 1 2 (MHNG); 1 2,
12.7.[?18...] (MHNG); 1 2, 15.7.1884, leg. H. Friese, coll. A. von Schulthess (ETHZ); 1 9,
6.9.1887, leg. M. Paul, coll. A. von Schulthess (ETHZ); 1 2, 18.7.1900, 1 2, 29.6.1901, 3 ©,
30.6.1901, leg./coll. E. Frey-Gessner (MHNG); 3 2, 26.6.1901, coll. A. von Schulthess
(ETHZ); 2 2, 5.7.1908, leg. T. Steck (NMBE). Simplon, Berisal: 1 2, 29.7.1939, leg./coll. A.
Nadig (ETHZ). Sion: 1 ©, 30.7.1892, leg./coll. T. Steck (NMBA); 1 2, 30.8.1892, leg. T.
Steck (NMBE). Sion, Château de la Soie: 2 9, 9.7.1968, an Euphorbia (MHNG); 1 ©,
13.7.1968, an Euphorbia (MHNG). St. German: 1 9, 23.7.1970, leg./coll. F. Amiet (NMBE).
St. Niklaus: 1 2, 15.7.1925, leg./coll. E. Däniker (NMBE). Stalden: 1 2, 30.7.1924, leg./coll.
T. Steck (NMBA); 2 2, 9.8.1933, leg. M. Naef (WL); 1 2, 7.7.1997, leg. G. Carron (GC [nicht
untersucht]). Visp-Stalden: 1 2, 31.7.1932, 1 2, 8.8.1933, 1 2, 21.7.1935, leg. M. Naef (WL).
Zeneggen: 1 9, 14.7.1970, 1 9, 23.7.1972, 1 2, 8.7.1973, leg. F. Amiet (FA); 1 2, 22.7.1986,
leg./coll. F. Amiet (NMBE).
FRANKREICH: Ariège, Verdun 1 d (SMNS); Korsika, Vizzavona 1 © (MHNG). GRIE-
CHENLAND: Hellas, Oros Killini, Ano Trikala 1 9 (WA). ITALIEN: Postumia 1 £ (DEI); Sizilien,
Taormina 2 d (ETHZ), 3 2,2 8 (ZSM); Vinschgau, Kastelbell 1 3 (ES); Vinschgau, Latsch 1
dé (ES); Vinschgau, Malserheide 1 SR (EIIHZ); Vinschgau, Schluderns 3 9, 3 d (BZOL).
KROATIEN: Krk, Baska 1 2 (ZSM). ÖSTERREICH: Burgenland, Piesting 1 £ (NMW); Bur-
genland, Winden 10 2 (NMW); Wien, Donau-Au | £ (BZOL). SLOWENIEN: Istrien, Portoroz
[Pirano] 1 3 (ZSM). SPANIEN: Leon, Ponferrada 1 £ (CS). TÜRKEI: Kayseri, Göreme 1 £ (CS);
Taurus, Marasch 1 9 (ZSM).
Diagnose ® ähnlich der von L. dorsigera, aber: Gelbe Zeichnung an den
Beinen meist leicht rötlich verfärbt; Pronotum vorne mit langem, hinten mit kurzem
Querband (Abb. 4a); Mesoscutum meist mit zwei Flecken im zentralen Bereich,
durchwegs mit zwei länglichen Flecken lateral (ähnlich L. gigas, Abb. 3d); Scutellum
am Hinterrand mit in der Mitte stark eingeschnürtem Querband; Metacoxa höchstens
ventro-lateral mit kleinem undeutlichen Apikalfleck; Femora apikal zu 1/2 bis 1/4
380 HANNES BAUR & FELIX AMIET
hell, Metafemur manchmal basal mit zusätzlichem Fleck (Abb. 2a"); Tibien und
Tarsen hell, nur die beiden ventralen Kiele der Metatibia proximal dunkel; Gastral-
terga 1, 4, 5 und 7 mit variablen Querbändern (Abb. 4c), auf den Terga | und 4
durchgehend, auf den beiden hinteren Terga in der Mitte unterbrochen; erstes Quer-
band in der Mitte stark eingeschniirt (Abb. 4b).
Clypeus kaum vorgezogen und mehr oder weniger verrundet (ahnlich L.
bifasciata, Abb. 5a‘); Flagellum stärker keulenförmig, Glieder 2-4 wenig kürzer als
breit oder quadratisch; Pronotum mit 2 Querkielen: einer im hinteren Viertel, der
andere fast unmittelbar entlang des Hinterrandes (Abb. 4a); Dorsellum stärker aus-
gerandet, mit 2 deutlichen Zähnen am Hinterrand; Gastraltergum 1 ohne Längsfurche
(Abb. 4b), diejenige des 5. Tergums tief, so dass Ovipositorscheide in Ruhelage
teilweise verdeckt ist (Abb. 4c); Ovipositorscheide kürzer, 1.02-1.23 mal so lang wie
der Metafemur (Abb. 12), reicht bis zum basalen Fünftel des 5. Tergums.
Höhenverbreitung n=64 50% = Phänologie n=69
0% 25% 50 % 75% Mai Juni Juli August September
ABB. 11, L. biguetina: Höhenverbreitung, Phänologie und Verbreitung in der Schweiz: @ Funde
nach 1950, O Funde vor 1951, potentielle Wirte (Auswertung von 15 Datensätzen von Osmia
acuticornis und O. tridentata).
LEUCOSPIDAE (HYMENOPTERA) DER SCHWEIZ 38]
Körperlänge 5.3-11.3 mm (Abb. 13), Kopfbreite 2.05-2.60 mm (Abb. 14).
Diagnose d: Gelbe Zeichnungselemente gelegentlich reduziert; Quer-
bänder auf dem Gaster meist durchgehend, Querband auf 1. Tergum nicht ein-
geschnürt.
Form des Gasters ähnlich L. dorsigera (Abb. 2b’).
Körperlänge 7.0-9.2 mm (Abb. 13), Kopfbreite 2.10-2.40 mm (Abb. 14).
Verbreitung: Paläarktisch, östlich bis Tadschikistan (Bouéek, 1974); in
Mitteleuropa bisher nur aus der Schweiz (z. B. Schletterer, 1890; Madl, 1990),
Tschechien, Slovakei (Bouéek, 1959) und Österreich (Madl, 1989) nachgewiesen. In
der Schweiz ist das Vorkommen auf das Wallis beschränkt (Abb. 11), wo L. biguetina
oft auch nach 1950 (57% der Daten, 69% der Fundpunkte auf der Verbreitungskarte)
gesammelt wurde. Bemerkenswert ist das Fehlen von Daten aus dem unteren Teil des
Wallis, d. h. unterhalb von Sion. Die Verbreitung stimmt recht gut mit derjenigen der
potentiellen Wirte überein, wobei diese in der Schweiz weiter verbreitet sind (Nord-
schweiz, GE, GR, TI) (Abb. 11).
Höhenverbreitung: von 500 bis 1600 m ti. M. (Abb. 11).
Gefährdung: gefährdet. Die beiden potentiellen Wirte (siehe unten)
werden in der Roten Liste (Amiet, 1994) für die Südschweiz als gefährdet aufgeführt.
Biologie: Aus Nestern von Osmia acuticornis Dufour & Perris (Le Goff,
1996, 1997b sub Hoplitis) und O. tridentata Dufour & Perris (Le Goff, 1997a, 1997b
sub Hoplitis) ın Frankreich gezogen.
Imagines erscheinen von Juni bis September (Abb. 11) und werden normaler-
weise an xerothermen Standorten wie Steppen- und Trockenrasen angetroffen. Ein-
zelne Exemplare konnten auf Blüten von Doldengewächsen (Apiaceae) und Euphor-
bia (Euphorbiaceae) beobachtet werden (vgl. Material; Amiet, Merz, pers. Beob.).
Nach Bouéek (1974) vermehrt sich L. biguetina in Mitteleuropa parthenogenetisch,
wobei aus unbefruchteten Eizellen fast durchwegs Weibchen entstehen (Thelytokie).
Dies trifft wahrscheinlich auch auf die Population im Wallıs zu, da im reichen
Untersuchungsmaterial von insgesamt 69 Exemplaren kein einziges Männchen vor-
handen war. Allerdings scheint es sich hier um ein lokales Phänomen zu handeln,
denn aus anderen, nahegelegenen Gebieten, z. B. dem Vinschgau im Südtirol,
befanden sich unter insgesamt 9 Belegexemplaren 6 Männchen (vgl. Material).
Leucospis bifasciata Klug, 1814 5a’, 5b', 12-14
Untersuchtes Material: ASERBEIDSCHAN: Lenikoran, Azfilial Girkan Reserve 1 & (MH):
NW Baku, Varafta Mts. 1 ¢ (MH). «Asia Minor: Namzur» 1 2 (ZSM). BULGARIEN: Slancev
Briag 1 © (BZOL); SW Melnik 1 4 (BZOL). FRANKREICH: Basses-Alpes, Valensole 1 £ (FA),
2 2 (NMBE). GRIECHENLAND: Kozani, Siatista, Brücke 1 £ (BZOL); Peloponnes, Zacharo
1 2(ZSM); Preveza SB, 2 km S Kerasona 2 d (BZOL). ISRAEL: Antipatris 1 2 (ETHZ).
JORDANIEN: N Shuna 1 4 (BZOL); Safi, 50 km of Karak 1 & (BZOL). ?LIBANON: «Sarepta»
1 2(MHNG). SYRIEN: 20 km NE Latakia 1 2 (BZOL). TÜRKEI: Sultan Daglari, Yalvat 1 ©
(BZOL). TURKMENIEN: 15 km N Aschapat 1 2, 1 d (BZOL); Kopet-Dag Kizi-Arvat, 50 km S
Chajagaia 1 2 (BZOL). USBEKISTAN: Yangikichlak, 10 km NW Ddjiak 1 d (BZOL).
Diagnose 9 ähnlich der von L. dorsigera, aber: Scutellum am Hinterrand mit
in der Mitte leicht eingeschniirtem Querband, dieses oft die Mitte des Scutellums
iiberschreitend; Querband auf Gastraltergum 5 in der Mitte unterbrochen.
382 HANNES BAUR & FELIX AMIET
'.n=13____n=88 n=14_ n=186 _n=61___n=123_ n=79_n=18
mig de (©)
16—
14=
1.2 ==
1.0-
©)
Ar
0.8—
I I TT I I ra
bifasciata brevicauda intermedia gigas Form 'A'
biguetina dorsigera gigas Wallis gigas Form 'B'
ABB. 12, Boxplot (nach Tukey) von Leucospis: Verhältnis Ovipositorscheide- zu Metafemur-
länge. Die Grafik zeigt Median, Interquartilbreite, Variationsbreite (ohne Ausreisser), Aus-
reisser (O), Extremwerte (%).
Clypeus kaum vorgezogen und mehr oder weniger verrundet (Abb. 5a');
Flagellum stärker keulenförmig, Glieder 2-4 wenig kürzer als breit oder quadratisch;
die beiden vorderen Querkiele des Pronotums kräftiger entwickelt als der hintere Kiel;
Längsfurche des 1. Gastraltergums an der Basis deutlich verjüngt (Abb. 5b’); Ovi-
positorscheide kürzer, 0.90-1.05 mal so lang wie der Metafemur (Abb. 12), reicht
höchstens bis zur Mitte des 1. Tergums.
Körperlänge 6.3-9.0 mm (Abb. 13), Kopfbreite 1.85-2.60 mm (Abb. 14).
Diagnose d: Gelbe Zeichnungselemente gelegentlich reduziert; Quer-
bänder auf dem Gaster normalerweise durchgehend.
Form des Gasters ähnlich L. dorsigera (Abb. 2b’).
LEUCOSPIDAE (HYMENOPTERA) DER SCHWEIZ 383
Weibchen
_n=13 __.n=88 __n=14_ _n=187 __n=61 n=123__ n=79 _
16=
14=
O
| O
12—
O
| o
10— MES È TE
PE
8— —
i | | |
(Sra did 0
6= na ce
O Ben
(©)
bifasciata brevicauda intermedia gigas Form 'A'
biguetina dorsigera gigas Wallis gigas Form 'B'
Mannchen
nn MENS, n26 1e NES6 n=10... n=42
14—
25,
| o
10—
wi
8— F E: I
A — A
| |
6= pela
bifasciata biguetina brevicauda dorsigera gigas intermedia
ABB. 13, Boxplot von Leucospis: Körperlänge in mm.
Körperlänge 4.7-8.3 mm (Abb. 13), Kopfbreite 1.40-2.30 mm (Abb. 14).
Verbreitung: Paläarktisch, östlich bis Tadschikistan (Bouëek, 1974); in Mittel-
europa wurde L. bifasciata bisher noch nicht nachgewiesen. Der Schweiz am nächsten
384 HANNES BAUR & FELIX AMIET
gelegene Vorkommen: Frankreich, Basses Alpes, Valensole (FA, NMBE); Italien,
Piemont, Oulx und Veneto, Venedig (Boucek, 1974).
Biologie: Aus Nest von Anthidium strigatum (Panzer) in Kroatien ge-
zogen (Fahringer, 1922); Bestimmung des Parasitoiden überprüft von Bouëek (1974).
Leucospis brevicauda Fabricius, 1804 2a-b, 12-14.
Untersuchtes Material: ALGERIEN: El Afroun 1 d (MHNG). ITALIEN: Puglia, Mt.
Gargano, Cagnavo 1 © (BZOL); Sardinien, Siniscola 1 £ (FNSM). MAROKKo: 4 2 (MHNG);
70 km N Agadir, Tamri 1 © (BZOL); 100 km E Bouzakame, Icht | d (BZOL); Tanger 4 9,
3 6 (MHNG); 10 km S Taroudant 1 4 (BZOL). TUNESIEN: Gabès, Skhira 2 2 (MH); Gafsa-El
Guetar 1 2 (BZOL); Metouia 1 d (MH).
Diagnose ® ähnlich der von L. dorsigera, aber: Flagellum rötlich;
Zeichnung neben schwarz und gelb auch rotbraun; gelbe Zeichnung häufig stärker
ausgedehnt als bei L. dorsigera, besonders an Pronotum, Mesoscutum, Scutellum und
Gaster; Metafemur ausser an den Zähnen und an der Basis gelb (Abb. 2a); Metatibia
dorsal rotbraun bis schwarz, höchstens in der Mitte mit feinem gelblichem Längs-
streifen; Tarsen meist rotbraun; Gastralterga 1, 4, 5 und 7 mit variablen Querbändern,
auf den Terga | und 4 manchmal, auf Terga 5 und 7 immer in der Mitte unterbrochen;
5. Tergum meist zusätzlich mit kleinen Flecken; auf dem 7. Tergum helle Zeichnung
rotbraun, daher oft undeutlich.
Clypeus kaum vorgezogen und mehr oder weniger verrundet (ähnlich L.
bifasciata, Abb. Sa'); die beiden vorderen Querkiele des Pronotums kräftiger ent-
wickelt als der hintere Kiel; basaler Zahn des Metafemurs ungefähr so lang wie
folgende Zähne (Abb. 2a); Gastralterga 1-4 ohne Längsfurche (ähnlich L. biguetina,
Abb. 4b), diejenige des 5. Tergums tief, so dass Ovipositorscheide in Ruhelage
teilweise verdeckt ist (ähnlich L. biguetina, Abb. 4c); Ovipositorscheide sehr kurz,
0.66-0.74 mal so lang wie der Metafemur (Abb. 12), reicht nur bis zur Mitte oder zum
basalen Drittel des 5. Tergums.
Körperlänge 5.8-9.2 mm (Abb. 13), Kopfbreite 1.90-2.45 mm (Abb. 14).
Diagnose d: Gelbe Zeichnungselemente gelegentlich reduziert;
Querbänder auf dem Gaster durchgehend.
Gaster 1m basalen Drittel stark verschmälert (Abb. 2b).
Körperlänge 6.8-8.2 mm (Abb. 13), Kopfbreite 1.45-2.15 mm (Abb. 14).
Verbreitung: Circum-mediterran (Bou,,ek, 1974); in Mitteleuropa
wurde L. brevicauda bisher noch nicht nachgewiesen. Der Schweiz am nächsten
gelegene Vorkommen: Frankreich, Hérault, Palavas (Berland, 1934b) und Bouches-
du-Rhöne, Port du Bouc (Bouëek, 1974): Italien, Puglia, Mt. Gargano (BZOL).
Biologie: Wirt unbekannt.
LEUCOSPIDAE (HYMENOPTERA) DER SCHWEIZ 385
Weibchen
__n=13 __n=88 _n=14 « n=187 n=61 n=123 n=79 n=18
4.0—
3.5= |
oi le —- ape
a
A
TH
IS Q
bifasciata brevicauda intermedia gigas Form 'A'
biguetina dorsigera gigas Wallis gigas Form 'B'
Männchen
Zn INT m eee Ai Ve EURE a
3.0— calli
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2.5= |
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2.0— Pre wa
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bifasciata Core Poe dorsigeran gigas intermedia
ABB. 14, Boxplot von Leucospis: Kopfbreite in mm.
386 HANNES BAUR & FELIX AMIET
DANKSAGUNG
An erster Stelle gebührt unser Dank Elsa Obrecht (NMBE) für die sorgfältige
Durchsicht des Manuskriptes sowie die Ausleihe des Materials am NMSO. Rene
Hoess, Bern, Schweiz, las eine frühere Version des Textes, und wir verdanken ihm
viele nützliche und kritische Anregungen. Wir danken ebenfalls einem anonymen
Rezensenten. Charles Huber (NMBE) hatte entscheidenden Anteil bei der Erstellung
der Verbreitungskarten, für die Geduld gegenüber den vielfältigen Wünschen des
einen Autors (H.B.) sei herzlich gedankt. Bertrand Baur, Rumendingen, Schweiz,
begleitete H.B. auf einer Exkursion ins Wallis und machte Habitataufnahmen. Ingmar
Wall (IW) überliess uns bereitwillig seine Notizen zu einem Bestimmungsschlüssel
der europäischen Leucospis-Arten. Frank Burger (FB), Jakob Forster, Winterthur,
Schweiz, Gerd Reder (GR), Albert Sermet (ASE), Erwin Steinmann (ES), Hansueli
Tinner-Guler (HT) und Jürg Zettel, Zoologisches Institut, Bern, Schweiz, teilten uns
ausführlich Freilandbeobachtungen zum Verhalten einiger Arten mit und stellten
zusätzlich eigenes Sammlungsmaterial bzw. hervorragende Dias zur Verfügung.
Ihnen allen sei herzlich gedankt. Für die Ausleihe von Material oder Angaben zu
Exemplaren in Sammlungen danken wir schliesslich den folgenden Personen und
Institutionen: Werner Arens (WA), Stefan M. Blank (DEI), Hermann Blöchlinger,
Naturmuseum Thurgau, Frauenfeld, Schweiz, Zdenék Boucek, The Natural History
Museum, London, UK, Daniel Burckhardt (NMBA), Toni Bürgin, Naturmuseum, St.
Gallen, Schweiz, Heinrich Bürgis (HB), Gilles Carron (GC), Erich Diller (ZSM),
Emanuel Gerber, Musée d’histoire naturelle, Fribourg, Schweiz, Fritz Gusenleitner
(BZOL), Martin Hauser (MH), Ulrike Hausl-Hofstätter, Landesmuseum Joanneum,
Graz, Österreich, Peter Huemer, Tiroler Landesmuseum Ferdinandeum, Innsbruck,
Österreich, Frank Koch (MNHU), Jens-Peter Kopelke (FNSF), Manfred Kraus,
Nürnberg, Deutschland, Walter Linsenmaier (WL), Ivan Löbl und Bernhard Merz
(MHNG), Paul Mildner, Landesmuseum für Kärnten, Klagenfurt, Österreich, Andreas
Müller (ETHZ), Till Osten (SMNS), Enrico Ratti (MSNV), Helmut Riemann,
Ubersee-Museum, Bremen, Deutschland, Ali Salvioni (AS), Michel Sartori (MZL),
Christian Schmid-Egger (CS), Stefan Schödl (NMW), Ingmar Wall.
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REVUE SUISSE DE ZOOLOGIE 107 (2): 389-418; juin 2000
The Bufo tuberosus species group with the description
of a new species from the rainforest of Cöte-d'Ivoire
Mills TANDY! & Jean-Luc PERRET?
! Colorado State University
201 Highland Drive
Sierra Vista, Arızona 85635 USA.
? Département d’herpétologie et d’ichtyologie
Muséum d’histoire naturelle
Route de Malagnou 1, case postale 6434
CH-1211 Geneve 6, Suisse.
The Bufo tuberosus species group with the description of a new species
from the rainforest of Cöte-d’Ivoire. - A very small species of toad
similar to Bufo tuberosus was discovered in a collection made in West
Africa by M. Lamotte in the late 1960s. In the 1980s a second specimen of
this species from Cöte-d’Ivoire was donated to the Muséum d’histoire
naturelle de Geneve. This paper describes the new species, B. amieti,
presents additional data on B. tuberosus, compares these species to others
and hypothesizes an origin of B. amieti and B. tuberosus by geographical
isolation of a common ancestor in eastern and western blocks of equatorial
rainforest separated by the V-Baoulé during past interpluvial periods.
Key-words: Bufo - Bufo tuberosus group - speciation - Céte-d’ Ivoire - rain
forest - West Africa - Anura.
INTRODUCTION
This paper summarizes the biology of the central African rainforest toad Bufo
tuberosus Günther and describes a new species of the same group from the far
western section of African equatorial lowland rainforest in Côte-d'Ivoire.
Bufo tuberosus, a very distinctive species from the rainforests of central
Africa, was described in the mid nineteenth century. It is so morphologically different
from other African Bufo with its usually globose parotoid glands and extreme
development of spinose warts that it was placed in its own species group (Tandy,
1972; Tandy & Keith, 1972). Although Bufo tuberosus was described very early in
the literature on African amphibians, and collections have been made by various
investigators over the years, it is not a common amphibian in museum holdings.
Little is known about its behavior and nothing about its genetics. This paper analyzes
Manuscript accepted 08.01.2000
390 MILLS TANDY & JEAN-LUC PERRET
morphological variation based on museum material, reports observations of its calling
behavior and recordings of ıts mating call made in Camerounian rainforests by Amiet
(1973, 1975, 1976, 1989) and by Tandy & Tandy (1976) and considers ecological
features of known localities.
In the 1970’s, Tandy encountered a small toad in a collection from Graba-
zouo, Côte-d'Ivoire made by M. Lamotte and R. Vuatroux. This is a gravid female
that resembles Bufo tuberosus but which is half the size of adult females of that
species. In 1986, J.-L. Perret received a second female of this species, collected by R.
Neumeyer, from Tai, Cöte-d’Ivoire. This paper describes these specimens as a new
species and compares them to Bufo tuberosus and other species from central and
western Africa. The new species is named in honor of our colleague who has contri-
buted so much to central African herpetology, J.-L. Amiet.
Terminology and methods of data collection and analysis are given in Tandy
(1972), Tandy & Keith (1972), Tandy & Tandy (1976), Largen et al. (1978), Tandy et
al. (1982), Tandy er al. (1985) and Tandy & Feener (1985).
Data are presented in the following format: locality, geographic coordinates;
sex and collection numbers of specimens; collection numbers of tape recordings; alti-
tude; date of collection; collector.
The following abbreviations are used for collection numbers:
AMNH American Museum of Natural History, New-York;
BM British Museum (Natural History), London;
CAS California Academy of Sciences, San Francisco;
CNHM Chicago Natural History Museum, Chicago;
FM Field Museum of Natural History, Chicago;
MBG Mission Biologique au Gabon;
MCZ Museum of Comparative Zoology,
Harvard University, Cambridge, Massachusetts;
MHNB Muséum d’histoire naturelle, Bale;
MHNG Muséum d’histoire naturelle, Geneve;
MHNCF Muséum d’histoire naturelle, La Chaux-de-Fonds;
MNHP Museum national d’histoire naturelle, Paris;
MT Collection of M. Tandy;
MT() Reference number of M. Tandy assigned to specimen of collection
indicated within parentheses;
MT.TC.Ca Collection of tape recordings of M. Tandy; tape cut from Cameroun;
RGMC Registre général du Musée du Congo, Tervuren;
SMF Natur-Museum und Forschungsintitut Senckenberg, Frankfurt;
UMMZ Museum of Vertebrate Zoology, University of Michigan, Ann Arbor,
Michigan;
ZFMK = Zoologisches Forschungsinstitut und Museum Alexander Koenig, Bonn;
ZMB Zoologisches Museum der Humboldt Universitat, Berlin.
Bufo amieti sp. n.
Holotype: 1 adult female (MHNG 2594.17) from lowland rainforest near Tai, Tai Na-
tional Park, Côte-d'Ivoire (5° 52° N 7°28’W, altitude 123m) collected in 1986 by R. Neumeyer.
THE BUFO TUBEROSUS SPECIES GROUP 39]
Fic. 1. Bufo amieti sp. n. Holotype female (above): habitus, natural size; (below): detail of the
enormous paratoid gland, tympanum and eye.
Paratype: 1 adult female (MNHP 1997 4934 = MT(P) 6141) from lowland rainforest
on the Sassandra River near Grabazouo, Soubré, Cöte-d’Ivoire (5° 50’N 6° 35°W, altitude
~150m) collected 10.XII.1967 by M. Lamotte and R. Vuatroux.
Diagnosis
A very small species of Bufo resembling Bufo tuberosus Giinther, 1858, but
only half the size of that species. Body measurements and their ratios indicate at least
five diagnostic differences with all species to which B. amieti has been compared
392 MILLS TANDY & JEAN-LUC PERRET
Fic. 2. Bufo amieti sp. n. Holotype female: dorsal and ventral color pattern. Magnification 2X.
THE BUFO TUBEROSUS SPECIES GROUP 393
Fic. 3. Bufo amieti Sp. n. Paratype female: dorsal and ventral color pattern. Magnification 2X.
394 MILLS TANDY & JEAN-LUC PERRET
Fic. 4. Bufo amieti sp. n. female (paratype: MHNP 1997 4934); snout-urostyle length 35.9 mm.
Grabazouo, Soubré. Côte d'Ivoire; Bufo tuberosus female (BM 1909.7.9.14); SU 65.8 mm.
Efangono, Cameroun; B. tuberosus male (BM 1934.12.1.14); SU 45.0 mm. Lomié District,
Cameroun.
(Tables | - 3). Comparisons have been made with all species of Bufo known from the
geographic area concerned, west and central Africa, and with all members of the Bufo
regularis Complex because of the acoustical similarities between the mating call of
Bufo tuberosus and species of that complex. The ratio of tympanum diameter/parotoid
width separates B. amieti from almost all congeners. Tympanum diameter, snout-
urostyle length or parotoid length/parotoid width distinguish it from the others.
Tympanum distinct, its horizontal diameter approximately 0.12 the width of the head
and 0.18 the width of the parotoid; snout-urostyle length about 37mm; parotoid length
1.7 times the parotoid width; no tarsal fold; male unknown; first finger longer than
second; toes not extensively webbed (33/4 phalanges of fourth toe free of webbing on
both inner and outer margins); lacking series of warts on posterior surface of forearm;
large warts with multiple cornified spinules extensively developed. Ecological data
for two localities: lowland rain forest, altitude,
X = 136.5m, range (123-150); mean annual rainfall,
x = 173.70cm (161.1-186.3); mean annual temperature, x = 23.95°C (23.9-24)(Table 4).
DESCRIPTION
Holotype: Snout-urostyle length (SU) 37.7mm; form moderately stout; head
triangular, broader than long, head width (HW) 15.5mm, head length (HL) 10.2mm;
snout obtusely rounded; nostrils closer to tip of snout than to eye; canthus rostralis
395
THE BUFO TUBEROSUS SPECIES GROUP
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TABLE 2. Apparent diagnostic differences between linear morphological characters of adult
females of Bufo amieti and species of Bufo from west and central Africa and those all species of
the B. regularis Comples. * species known from west of the Dahomey Gap. These comparisons
are tentative because of the small sample size for B. amieti (n=2). A difference (+) indicates
that ranges of variation do not overlap. These comparisons cannot be tested for statistical
significance because of the small sample size for B. amieti and a few of the other species. For
data on other species, see Table 7 in Largen er al. (1978), Table 12 in Tandy et al. (1982) and
Tables 1, 9 and 16 in Tandy & Feener (1985).
© ©
= =) > 2 7
| ed i |
Species DE à RME an 2 32
Sex py yar AE NS EN ac ON ec ec CE | ws SAS
Females
B. tuberosus er E ati + + + 8 9
B. gracilipes + + + + ++ + 7 10
B. villiersi + + + + + + + + + + + # 12 5
B. regularis* + + + + + + + + + + + 11 6
BaSUDENCIILONIS mcr haan ASSE EER ts + + + + + + 14 3
B. garmani + + + + + + + + + + + 11 6
B. latifrons + + + + + + E EE nl 6
B. togoensis* IB. Sp) geo! ae Dar Ar + + + + + 112 5
B. brauni + + + + + + + + + + + + + + 14 3
B. gutturalis Fall Epto. Peso = + + ee a in 6
B. kisoloensis + + + + + + + + SPE + 12 5
B. poweri È SP Wt. Apa enon ae AR. ara dr ESEL u} 14 3
B. rangeri Ai ap, Gp SP Ge ir + + © 10 7
B. camerunensis* + + + + + + + + + E LS tk 13 4
B. kerinyagae + + + + + + + qi)
B. xeros* A A e NUE A + PAE ar a ae 12 5
B. asmarae + + + + + + - + + + + + 12 5
B. maculatus* papi CES Ea a - + + + + + + 13 4
B. danielae* + + + + + - + + + 9 8
B. blanfordi + 4 + IG: > 12
B. langanoensis + + + + + + O Hi
B. turkanae + + + + + + + + + + + + + 13 4
B. perreti CO PAU A SEA IO. - + Su tah fe hd Sb 13 4
B. steindachneri + + Sey CE eos 8 9
B. pentoni*
- Senegal + + + + + Si + + +
+
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N
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in relation to body size, globose, 1.6 times longer than wide, (PL/PW 1.60), narrowly
separated from the eye, the anterior edges lying between the anterior and posterior
borders of the tympanum, parotoid length (PL) 9.6mm, 26% of snout-urostyle length,
parotoid width (PW) 6.0mm, 63% of parotoid length.
First finger longer than second, second longer than fourth; fingers lacking
marginal folds; subarticular tubercles large, undivided; palm with numerous tubercles
lacking spinules; inner metacarpal tubercle moderately developed but much smaller
than outer metacarpal tubercle and lacking spinules; lacking any series of distinct
warts on the posterior surface of the forearm; toes moderately webbed; toe IV with
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402 MILLS TANDY & JEAN-LUC PERRET
with only a slight margin of webbing at the base; webbing with small melanized
spinules on the dorsal surface; tubercles of soles smaller than those of palms; inner
and outer metatarsal tubercles prominent and suited for burrowing, the inner larger
and more raised than the outer. No tarsal fold. Urostyle-heel length (UH) 26.5mm.
Dorsum with numerous conical and rounded warts, each with one or more large
spinules and some smaller spinules; spinules extending over entire dorsum, including
surfaces of parotoid glands, but less numerous anterior to eyes. Venter with somewhat
smaller warts with spinules similar to those of the dorsum. Warts at rictus partially
fused and covered with numerous small spinules.
Ovaries with immature eggs. The specimen was apparently collected outside of
the breeding season.
Color (in alcohol) disruptively patterned light and dark brown; Dorsal back-
ground color light brown; four pairs of bilateral reticulate dark brown markings. No
B. regularis-like white spots on dorsum; No vertebral line; Parotoids and rictal warts
same color as dorsum; Dark melanized spinules prominent against lighter back-
ground. Lower margin of orbit and part of area beneath eye cream. Upper surfaces of
limbs with well defined brown cross bars sometimes outlined in darker brown;
posterior femoral integument light brown. Venter markedly reticulately mottled
brown and cream. See Figs. | - 2.
Paratype: The paratype female is similar to the holotype (Table 1). It is slightly
smaller- SU 35.9mm, but is gravid and thus indicates that the breeding season
includes December near the beginning of the long dry season. Coloration is like the
holotype except overall a somewhat darker brown, and it has a few B. regularis-like
white spots on the dorsum. See Figs. 3-4.
GEOGRAPHIC DISTRIBUTION AND ECOLOGY
B. amieti is known from only two localities in lowland rainforest of western
Cöte-d’Ivoire (Fig. 5). Statistics on the altitude, rainfall and temperature of these
localities are given in the diagnosis and in Table 4.
The small number of known localities for B. amieti precludes statistical
comparisons of ecological variables with other species. Data in Table 4 and 5 indicate
some differences and similarities. B. amieti seems to differ in one or more ecological
characteristics from eighteen of the 25 species with which it has been compared
(Table 5), and it is unlikely to be sympatric with any of those. Of the remaining seven
species, four, B. regularis, B. superciliaris, B. camerunensis and B. maculatus, are
known from west Africa west of the Dahomey Gap, but B. amieti has not yet been
found syntopic with any other bufonid species. Bufo cristiglans Inger & Menzies,
1961, is considered a synonym of B. camerunensis Parker, 1936.
If B. amieti is restricted to relatively undisturbed lowland rainforest, it is
unlikely to occur in the same local areas as B. regularis or B. maculatus. But such a
preference for high forest would make it more likely to occur with B. superciliaris or
B. camerunensis. If B. amieti does occur locally with these species, it would probably
avoid them because the small size of B. amieti would make it likely food fare for the
other two species.
403
UP
EROSUS SPECIES GRO
THE BUFO TUB
‘SASO12qN] ‘ = Soi) uodo ‘u ‘ds yonum 'g
= SSOI9 YIM SIMI SHSO12QM 'g pue 1
onup ofng JO UONNALISIP ILyde13093 UMOUY ‘G ‘DIH
0 M 0%
404 MILLS TANDY & JEAN-LUC PERRET
TABLE 4. Ecological characteristics of Bufo amieti and other species of Bufo that are known to
occur in west or central Africa. Accuracies and sources of these data vary. Some figures are
based on surveys or meteorological stations near the locality. Others were interpolated from
map contours, isotherms or isohyets. One decimal point given does not necessarily imply such
accuracy of measurement but rather that these were the figures used to compute the statistics.
Individual values for some altitudes may be in error by 50m or more. Most basic data are from
Bartholomew (1985), Knoch & Schulze (1956), NIMA (1999), Wernstedt (1959, 1972) or
Survey of Kenya (1962).
n Elevation Mean Mean
meters Annual Annual
Temp oC Rainfall cm
Bufo amieti
Cöte d’Ivoire
Grabazouo 150 23.9 161.1
Tai 123 24 186.3
Mean 2 136.5 23.95 1737
Bufo tuberosus
Cameroun, Gabon, Guinea Ecuatorial, République Centrafricaine, République du Congo, République Démo-
cratique du Congo
Mean 412.8 23.69 224.87
Standard Error 46.63 0.316 20.472
Minimum 10 22 150
Maximum 1266 26.4 411.8
n 48 14 15
Bufo danielae
Côte d'Ivoire
Monogaga ] 10 25 150
Bufo camerunensis
Cameroun, Gabon, Nigeria, République du Congo & République Démocratique du Congo
Mean 10 526.9 23.1 184.4
Standard Error 95.49 0.49 13.92
Minimum 100 20 140
Maximum 1066.8 24.7 290
Bufo togoensis
Togo 3 447,3 22.8 141
Minimum 350 22.8 139.7
Maximum 589.8 22.8 142.3
n 3 I 2
Bufo latifrons
Cameroun & Guinea Ecuatorial
Mean 5 549.7 24.1 250.9
Minimum 50 22.4 145.3
Maximum 876.9 232 350
Bufo maculatus
Angola, Bissao, Cameroun, Ethiopia, Kenya, Liberia, Mozambique, Republique Centrafricaine, South Africa,
Zimbabwe
Mean 10 480.5 231 163.4
Standard Error 206.62 1.09 44.4
Minimum 19.5 26.7 491.2
Bufo superciliaris
Cameroun, Ghana, Republique Democratique du Congo
Mean 10 561.1 22.9 193.9
Standard Error 113.59 0.55 123,112
Minimum 50 20 150
Maximum 1066.8 26 260
Bufo regularis
Algeria, Angola, Cameroun, Egypt, Ethiopia, Gabon, Kenya,
République Démocratique du Congo, Senegal, Sudan, Uganda
THE BUFO TUBEROSUS SPECIES GROUP
405
Elevation Mean Mean
meters Annual Annual
Temp oC Rainfall cm
Libya, Nigeria, République Centrafricaine,
Mean 793.6 2117, 102.67
Standard Error 81.16 0.453 8.373
Minimum 7 14 0.3
Maximum 2500 29.6 280
Bufo xeros
Chad, Ethiopia, Kenya, Uganda
Mean 802.6 23.4 57.8
Standard Error 164.72 1.41 7.8
Minimum 54.9 16 29.9
Maximum 1676.4 29.2 100
Bufo pentoni
Haute Volta, Senegal, Sudan
Mean 131.5 DIET; 40.8
Minimum 5 25 14.9
Maximum 450 29 70
Bufo gracilipes
Cameroun, Gabon, Guinea Ecuatorial, Republique du Congo
Mean 310.81 2392 214
Standard Error 97.57163 0.345704 19.95348
Minimum 10 23 166.4
Maximum 780 23] 350
Bufo villiersi
Cameroun
Mean 1853.5 23.38 263.62
Minimum 1398 22.0 187.7
Maximum 2500 25.0 305.0
B. xeros and B. pentoni occur in much drier habitats in the northern portion of
West Africa.
B. danielae is known from only two coastal sites in Cöte-d’Ivoire (Perret,
1977) which are at lower elevation and appear to have somewhat warmer and drier
climates than do the known localities of B. amieti.
B. togoensis is known from only three localities that are at higher elevations
with somewhat cooler temperatures and lower rainfall than the known sites for B.
amieti. Tandy (1972) considered B. togoensis to be a synonym of B. latifrons. Addi-
tional recent collections indicate that B. togoensis is a distinct species resembling
B. latifrons in the size and proportions of its tympani and
B. camerunensis in the structure of its parotoid glands.
The ecological features of B. tuberosus localities in central Africa do not
appear to differ from B. amieti sites.
The similarities of both morphology and ecology indicate a close relationship
between B. amieti and B. tuberosus. Apparently B. amieti is a western isolate derived
from a common ancestor of the two species. This pattern is similar to that of several
pairs of lowland rainforest hyperoliid treefrog species east and west of the V-Baoulé
406 MILLS TANDY & JEAN-LUC PERRET
TABLE 5. Apparent differences and similarities between the ecological characteristics of Bufo
amieti and species of Bufo from west or central Africa and those of all species of the B.
regularis Complex. * species known from west of the Dahomey Gap. These comparisons are
tentative because of the small number of known localities for B. amieti (n=2). A difference (+)
indicates that ranges of variation do not overlap. These comparisons cannot be tested for statis-
tical significance because of the small sample size for B. amieti and a few of the other species.
(1) - species for which number of localities is less than 10. For data on other species, see Tables
3 in Tandy et al. (1976) and Tandy er al. (1982), Table 8 in Tandy er al. (1985), and Table 13
in Tandy & Feener (1985).
Species compared Elevation Mean Annual Mean Annual Different Similar
Temperature Rainfall
B. tuberosus = & 0 3
B. gracilipes - = 2 0 3
B. villiersi (1) + È in 2 1
B. regularis* - if 0 3
B. superciliaris* 0 3
B. garmani = = de I 2
B. latifrons (1) - È 0 3
B. togoensis*(1) + + fe 3 0
B. brauni (1) + di St 3 0
B. gutturalis - = At 1 2
B. kisoloensis + + 2 1
B. poweri + + fe 3 0
B. rangeri = er | 2
B. camerunensis* - 0 3
B. kerinyagae chi + È 2 1
B. xeros* An I 2
B. asmarae + + DI 1
B. maculatus* = A 0 3
B. danielae* (1) + + ane 3 0
B. blanfordi + = ae 2 1
B. langanoensis (1) + + N 3 0
B. turkanae (1) + SÈ n 3 0
B. perreti (1) + + È DI 1
B. steindachneri 2 i 1 2
B. pentoni* - Senegal (1) - + + 2 1
ecological break of Cöte-d’Ivoire as described by Schigtz (1967) for Hyperolius
sylvaticus ivoiriensis vs H. s. sylvaticus, Leptopelis macrotis vs L. rufus (erroneously
as “L. palmatus” (Perret, 1973)) and Leptopelis occidentalis vs L. boulengeri. The
initial evolutionary barrier to eastern and western isolates probably is related to
changes in Africa’s climate during geologic time and particularly to interpluvial
periods when the rainforest blocks were probably most isolated.
Bufo tuberosus Günther, 1858
Bufo tuberosus Ginther, 1858: 60, pl. 3. fig. C, 140. Peters, 1875: 202. Boulenger,
1880: 546-547, 564, 572-573; 1882: 304. Rochebrune, 1884: 20, 47-49, Pl. IV, Fig. 2.
Boulenger, 1887: 565. Bocage, 1895: 15, 272. Werner, 1898: 202. Mocquard, 1903: 215.
Boulenger, 1900: 435; 1903: 62. Bocage, 1903: 45. Boulenger, 1906: 158. Johnston, 1908:
950. Nieden, 1908: 509-510, 517. Muller, 1910: 625. Nieden, 1910: 66, 68, Fig 147. Barbour,
1911: 135. Lampe, 1911: 220. Arldt, 1917: 123. Noble, 1922: 39. Nieden, 1923:104. Noble,
1924: 167, 177-178, 311. De Witte, 1930: 241-242, 252; 1934: 167. Loveridge, 1936: 84.
Parker, 1936: 155. Mertens, 1938:10; 1941: 277. Monard, 1951: 174. Curry Lindahl, 1956: 56.
Perret & Mertens, 1957: 555, 557. Guibé & Lamotte, 1958: 243. Inger & Menzies, 1961: 594.
THE BUFO TUBEROSUS SPECIES GROUP 407
Mertens, 1965: 14.19. Oates, 1965:89. Perret, 1966: 309-311, 314-315. Amiet & Perret, 1969:
120. Perret, 1971: 131. Perret & Amiet, 1971: 48, 50, 52-53. Amiet, 1973: 136; 1975: 46, 50,
54, 56 (photo); 1976: 150-152 (sonagram); 1978:199. Frost, 1985:63. Amiet, 1989: 92, 95, 98-
99 (PI. I - photo). Brauer, 1991:6, 159, 179.
Bufo polycerus Werner, 1897: 211. Nieden, 1923: 105.
CAMEROUN - Akak-Yemefek; 2°37'N 10°04'E, alt. 50m; 2 MHNG917.67; 15.V.1955; J.-L.
Perret. Batouri; 4°35'N 14°24'E, alt. 655m; 6 BM1934.12.1.14; IV.1933; M. Merfield. Bibundi
(Bibunde); 4°13'N 8°59'E, alt. 50m; (Lampe, 1911). Bipindi; 3°06'N 10°30'E, alt. 200m; 1
specimen; Zenker. (Nieden, 1910). Campo; 2°22'N 9°48'E, alt. 10m; 2 MHNCF148; 1.1947;
A. Monard. Ebonji; 4°06'N 9°24'E, alt. 50m; "cri de rivalité" sonagram; (Amiet, 1989, p. 151).
Efangono (=?"Efayong"); 3°04'N 11°49'E, alt. 750m; 9 BM1909.7.9.14; 1908?; G. L. Bates.
Efulen, Kribi; 2°47'N 10°32'E, alt. 500m; d CNHM3585; Date? ; G. L. Bates. 2 BM1904.
10.26.28; 1903? ; G. L. Bates. 9 MHNB2482; 1906; Rosenberg. Ekekam III; 3°53'N 11°22'E,
alt. 750m; 6 MHNG1 176.5; 10.X.1968. 1 specimen; VI.1968. 3 specimens; X.1968-1.1969. J.
-L. Amiet. Foulassi; 2°59'N 11°58'E, alt. 650m; 2 d& MHNG1008.30-31; 2 MHNG917.66;
IV.1953; 49 2 MHNG1008.27-29, 1020.47; 8 im. MHNG1008.31-37, 1020.48-49; 1960-61; J.
-L. Perret. Gadji; 4°29'N 14°03'E, alt. 600m; ?BM1937.1.1.21; 1936?; M. Merfield. Idenao;
4°16'N 8°56'E, alt. 100m; 22 £ BM1968.470, -.508; 12.VIII.1967; T. Struhsaker. Iemonkonn;
3°54'N 11°58'E, alt. 700m; 1 specimen; I-II/1968. J.-L. Amiet. Kala; 3(51'N 11(22'E, alt. 350m;
recording of mating call; 12.1.1972; 2 specimens; VII-X.1967; 2 specimens; 15.II-15.1V.1969.
J. -L. Amiet. Kendonge (Kidonge), S. Bakundu Forest Reserve; 4°33'N 9°26'E, alt. 50m; d
BM1969.1635; 2 BM1969.509; 1968?; S. Gartlan. Kribi (includes "Dja River" of MCZ
record); 2°56'N 9°56'E, alt. 43m; 2 MCZ2660; 1908; G. H. Schwab. 39 2 UMMZ 35611; I-
VI.1907; 38320; 1908; 56277; no date; G. H. Schwab. Kumba (=Johann-Albrechtshöhe);
4°38'N 9°25'E, alt. 200m; 2 specimens; Conradt. Nieden, 1910). Limbe (Victoria); 4°01'N
9°12'E, alt. 100m; 4 specimens ZMB?13915; Preuss. (Nieden, 1908, 1910). 32 km N Lolodorf;
3°26'N 10°42'E, alt. 250m; d CAS103353 (TP3545); 21-22.III.1966; T. Papenfuss. Lomié
District; 3°09'N 13°35'E, alt. 632m; 2 BM1937.12.1.54; 1936?; M. Merfield. Mabiogo;
2°12'N 9°53 E, alt. 10m; 2 MHNCF88; 28.11.1947; A. Monard. Mbanlam; 3°17'N 9°54'E, alt.
612m; 6 MHNG955.100; 27.IX.1957; J. -L. Perret. Momobelenga; 3°56'N 11°40'E, alt. 600m;
3 sonagram of mating call; Date?; (Amiet, 1976, p.151). Mundame; 4°35'N, 9°31'E, alt. 100m;
2 specimens; 1906; Rohde. (Müller, 1910). Ngam/Sangmelima; 2°47'N, 11°54'E, alt. 759m;
3 22 MHNG955.97-99; 4.1.1957, 3.11.1957, 20.IX.1957; 1 specimen SMF52335; 17.VI.
1955; J. -L. Perret. Nkolfep; 3°59'N 11°24'E, alt. 350m; 1 specimen; 11.1968; J. -L. Amiet.
Nta Ali; 5°33'N 9°30'E, alt. 1266m; 3; 15.1V.1977; J. -L. Amiet (Amiet, 1978). Rio del Rey;
4°44'N 8°37E, alt. 50m; 1886?; H. H. Johnston. (Bouleng-er, 1887). Sangmelima (includes
"Zima Country" of BM records); 2°56'N 11°59'E, alt. 782m; 29 2 BM1906.5.28.150-151;
1905?; G. L. Bates. d CNHM19918; 1899; A. I. Good. nr Ting (32 km S Akonolinga) Sta.
141; 3°40'N 12°10'E, alt. 750m; record-ing of mating call, 4 MT.TC.CA123; 7.IV.1971. M.
& J. Tandy. Yapoma (nr Douala); 4°02'N 9°49'E, alt. 50m; 9 ZMB27541; 1910; Schafer. No
locality; ¢ ZMB8319; 1870-1880; A. Reichnow.
GABON - Makokou; 0°34'N 12°52'E, alt. 516m; 6 MHNG2207.33 (MBG1117); 25.X.1964; 9
MHNG2207.34 (MBG1277); 26.X1.1964; L. P. Knoepffler. Nzing-Ayong (=?Nzibelong);
0°18'N, 9°50'E, alt. 50m; 26 d MHNG2207.31-32 (MBG129, 336); 2 MHNG2207.30
(MBG128); 26.11.1964; L. P. Knoepffler.
GUINEA ECUATORIAL - Bioko (Fernando Poo): Basilé (Bassilé), slopes of Pico de Basilé (Santa
Isabel); 3°36'N 8°45'E, alt. 527m; 2 adults MB; 1894?; F. Newton. (Bocage, 1895, 1903-1905).
Rio Iladyi Falls, south of Bioko (Moca); 3°20'N 8°40'E, alt. 914m; 2 BM1969.2447;
18.1X.1964; University College Expedition. Luba (Bahia de San Carlos); 3°27'N 8°33'E, alt.
400m; im. SMF?; 20.IX-24.X.1962; R. Mertens. (Mertens, 1965). Moca (Moka); 3°20'N
8°40'E, alt. 914m; ZMB333999; XI.1939; H. Eidmann. (Mertens, 1941). Musola; 3°26'N
8°37'E, alt. 500-800m; (Boulenger, 1905-06). San Antonio de Ureca; 3°16'N 8°39'E, alt. 10m;
?ZFMK9291; 1.1963; M. Eisentraut. No specific locality, 9 BM1947.2.21.14(=BM1851-
10.25.7) Holotype; 1850; Fraser?
408 MILLS TANDY & JEAN-LUC PERRET
Rio Muni: Cabo San Juan (Cape St. John); 1°15'N 9°30'E, alt. 50m; Martinez de la
Escalera. (Boulenger, 1903). Macomo (Makomo); 1°43'N 9°49'E, alt. 350m. (Nieden, 1908).
Rio Uoro-Rio Mbini (Rio Benito); 1°34'N 9°38'E, alt. 50m; 4 2 9 BM1900.2.17.109-112;
1899; G. L. Bates. Rio Lana 2.4km N Evinayong; 1°39'N 10°39'E, alt. 750m; ?BM1965.1412;
6.1X.1965; J. Oates. No locality; ZMB20035; Date?; Collector?
REPUBLIQUE DU CONGO (Congo-Brazzaville) - Djamba; 0°05'S 15°43'E, alt. 350m; RG3159;
XII.1924; H. Schouteden. (De Witte, 1930).
REPUBLIQUE DEMOCRATIQUE DU CONGO (Belgian Congo; Zaire) - Bafwabaka; 2°10'N 27°50'E,
alt. 750m; 2 AMNH8405; 1909-1915; Congo Expedition. Buta; 2°47'N 24°47'E, alt. 350m; 2
specimens RG4402-03; 11.1925; G. F. De Witte. Epulu; 2°15'N 29°15'E, alt. 350m; 2 speci-
mens; 9.1V.1952. K. Curry-Lindahl. (Curry-Lindahl, 1956). Kifuku (s/la Nawa), ‚alt. ;
RG1595; VIII.1925; H. Schouteden. Medje; 2°25'N 27°18'E, alt. 750m; RG1054; IV-V.1910;
Lang & Chapin. Medje; 2°25'N 27°30'E, alt. 750m; AMNH8404; IV-V.1910; Congo Expedi-
tion. (Noble, 1924). Ngayu; 1°40'N 27°40'E, alt. 750m; AMNH8401-02; IV-V.1910; Congo
Expedi-tion. (Noble, 1924). (Note: AMNH8403 is a d Bufo funereus.)
REPUBLIQUE CENTRAFRICAINE - Tibili (=?"Tuburi, Narob"); 4°58'N 14°42'E, alt. 750m; 2BM
1912.1.11.19; 1911?; P. A. Talbot.
MORPHOLOGY
Many aspects of the morphology of B. tuberosus have been described and
illustrated in the literature. Illustrations include drawings in Giinther (1858), Roche-
brune (1884) and Nieden (1910) and photographs of preserved material in Perret &
Amiet (1971) and of living specimens in Perret (1966) and Amiet (1975, 1989).
PLATE 1. Bufo tuberosus female, Ngam, Sangmelima, Cameroun.
THE BUFO TUBEROSUS SPECIES GROUP 409
TABLE 6. Some diagnostic differences between linear morphological characters of adults of
Bufo tuberosus and species of Bufo from west and central Africa and those all species of the B.
regularis Complex. * species known from west of the Dahomey Gap. A difference (+) indicates
that ranges of variation do not overlap. Lack of a diagnostic difference does not indicate lack of
a statistical difference. These comparisons have not been tested for statistical significance. For
data on other species, see Table 7 in Largen er al. (1978), Table 12 in Tandy et al. (1982) and
Tables 1, 9 and 16 in Tandy & Feener (1985).
Species 3 =
Sex = = EME = A = = 5 5, £
ARE, ORE:
B. amieti* tp Bars ap En + + + 8
B. gracilipes + + + ER Credit 6 1
B. villiersi 0 17
B. regularis* + 1 17
B. superciliaris* + + + + + + + + + 9 8
B. gammani + + 2:
B. latifrons + + DIST;
B. togoensis* + + DNS
B. brauni + + + + + + + 8 9
B. gutturalis + + + Sun HA
B. kisoloensis + 1 16
B. poweri + + + + + + 6 1
B. rangeri + + + + + 5 12
B. camerunensis* + 1 16
B. kerinyagae + HTC + 4 13
B. xeros* + + + + + + + if 10
B. asmarae + + + + + + 6 1
B. maculatus* + 1 16
B. danielae* + + + + 4 13
B. blanfordi + + DIS
B. langanoensis + + 3 14
B. turkanae + + + + + + 6 11
B. perreti + + + + Ay WS)
B. steindachneri + + + + + + + 7 10
B. pentoni* - Senegal + + ZU
Males
B. amieti* a TAN ERNEST Ce oe a n) ? ?
B. gracilipes + + 2 IS
B. villiersi et + + 5) 12
B. regularis* + + Zils
B. superciliaris* + + + + + + + + + + + + 12 5
B. garmani See MORRIS al SEHEN RE + + + 9 8
B. latifrons + 1 16
B. togoensis* + + 2 15
B. brauni + + + + + + + + + 9 8
B. gutturalis + He + ++ + + ++ 10 7
B. kisoloensis + + + + + + Ol
B. poweri + + + + + + + + + + + 11 6
B. rangeri 2 A RE D INI SCH + + + + 10 7
B. camerunensis* + + 2 15
B. kerinyagae + + + + 5 12
B. xeros* + + + + + + + fetal
B. asmarae + + + + + 5 12
B. naculatus* + 1 16
B. danielae* + l 16
B. blanfordi + 1 16
B. langanoensis + + 3 14
B. turkanae + + + 3 14
B. perreti + + DOS
B. setindachneri + + 3 14
B. pentoni* - Senegal + + 2 AS
410 MILLS TANDY & JEAN-LUC PERRET
Günther’s description focused on the extreme wartiness of the skin, but did not
give measurements or ratios of body parts. The sex of the holotype was not specified
nor was its exact locality on the island of Bioko (Fernando Poo) (Günther, 1858).
Boulenger later published a more detailed description with some measurements and
verbal proportions, however he mistakenly identifed the holotype as a male and his
measurements for that specimen differ from ones made more recently.
The very prominent parotoids were noted by Boulenger (1880) and by sub-
sequent authors such as Noble (1924), Perret (1966) and Perret & Amiet (1971). The
data in Tables | and 6 demonstrate the markedly low ratio of parotoid length to width
and also the very small size of the tympanum compared to parotoid width. The ratio
of T/PW separates B. tuberosus diagnostically (ranges of variation do not overlap)
from 18 (in females) or 21 (in males) of the 25 other African Bufo species compared.
As summarized in Table 6, B. tuberosus differs diagnostically in at least one of these
measurements or their ratios from all of the species compared except for females of
B. villiersi.
Boulenger (1900) mentioned a crimson vertebral line in some specimens from
the Rio Uoro (Rio Benito), Rio Muni, Guinea Ecuatorial. That color has now faded in
those specimens.
Noble (1924) noted varıation in ventral coloration in his material from the
eastern République Démocratique du Congo ranging from heavy to no dark mottling.
Parker (1936b) gave ratios of body to foot length and interpreted this as
demonstrating relatively short legs of B. tuberosus compared to other species. Perret
(1966) gave ranges of variation of body and tibia lengths for each sex and ratios of
parotoid gland length/width and body length/tibia length. These data do not appear to
support Parker’s appraisal of relative leg length, although Perret did not give arith-
metic means of his ratios (only ranges) nor did he test such for statistical significance.
Although differences of means given in Table | have not been tested for significance,
the data on ratios of snout-urostyle length/urostyle heel length do not support Parker’s
judgement about leg length in this species. Those data indicate that B. tuberosus is
one of the longer legged species.
Perret (1966) noted the marked sexual dimorphism of body size in this species.
Tables 1 and 7 illustrate this dimorphism in linear characters. All measurements are at
least statistically smaller in males than females, and two are diagnostically different.
Most ratios do not differ between the sexes, although females have statistically
narrower heads relative to body size (SU/HW) and smaller tympani relative to paro-
toid width (T/PW).
TABLE 7. Sexual dimorphism of linear morphological characters in Bufo tuberosus. D -
difference diagnostic, ranges of variation do not overlap. S - statistically significant difference
P < 0.01. - no statistical difference
Number of characters
2 3 o
= = 3 Bo rà don mer
Boe 2 e a 2 È S BE Sl 2 ei) a à 2
»D S$ SS SS SH S S CR Ne 7)
THE BUFO TUBEROSUS SPECIES GROUP 411
Perret (1966) stated that he had not found vocal sac openings in males except
for one with a single opening on the left. Perret & Amiet (1971) made a similar
statement. Table 3 shows numbers and locations of vocal sac openings in the eight
adult males examined. Two lack openings.
MATING CALL
Basic physical structure. The mating call of Bufo tuberosus is a series of low
pitched hooting sounds that lack the noisy quality of the calls of most African Bufo.
They are also not as loud as those of many other species (Amiet, 1976). The physical
structure of these calls is similar to that of species of the B. regularis, B. maculatus, B.
blanfordi, B. funereus (except B. villiersi), and B. pentoni groups and also some
species of the B. vertebralis Complex and B. angusticeps group (Tandy, 1972; Tandy
& Keith, 1972; Tandy & Tandy, 1976). Each complex pulse train (series of hoots)
contains a variable number of simple pulse trains repeated at a rate of about 1.6 per
second in specimens from Cameroun (Table 8). Each simple pulse train contains
about 17 passive pulses repeated at a rate of 87 per second (89.74/sec at about 24°C).
The duration of simple pulse trains is about 0.2 second (.18 at 24°C). The low
emphasized frequency is the dominant frequency at about 526Hz. Sonagrams show
three harmonics above the low emphasized frequency, but these contain less than half
the energy of the low emphasized frequency. These frequencies show a slight gradual
rise at the ends of simple pulse trains. These data are based on calls of only two
recorded specimens. See material studied for times and places.
Amiet (1976) described and illustrated a sonagram of a call of a male B.
tuberosus uttered while trying to clasp a plastic bag containing another male which
Amiet termed a “cri de rivalité”. This call is structurally very similar to male release
calls uttered by many Bufo species, and the context indicates that is what it was. It is
very different from the mating call in structure and function.
TABLE 8. Physical characteristics of the mating calls of Bufo tuberosus. PPR = passive pulse
repetition rate; NPP/PT = number of passive pulses per simple pulse train; PT DUR = simple
pulse train duration; PTR = simple pulse train repetition rate; LOW EMP = low emphasized
frequency; DOM = dominant frequency; n = number of individuals per sample.
N PPR NPP/PT PT DUR PTR LOW EMP DOM
(/sec) (/sec) (/sec) (Hz) (Hz)
Bufo tuberosus
Kala, Cameroun 1
84.80 18.60 0.230 1.30 547.0 547.0
Sta. 141 nr Ting, 1
Cameroun 89.74 15.96 0.178 1.92 505.0 505.0
Circadian and seasonal calling behavior. In April, 1971, Tandy found a mixed
chorus of Bufo gracilipes and B. tuberosus calling along a small stream at dusk in
secondary forest near Ting, Cameroun. The B. gracilipes called in water holding onto
living aquatic plants and vegetative debris in the middle of the stream. B. tuberosus
called from very well-concealed sites among overhanging roots of forest vegetation
beneath the undercut stream bank. Amiet (1973, 1976, 1989) was able to make
412 MILLS TANDY & JEAN-LUC PERRET
additional observations of this behavior. The above two species and B. villiersi are the
only Camerounian Bufo known to typically call during daylight, and these species are
atypical of most species in the genus.
Amiet (1976) has found that B. tuberosus choruses are usually small, often less
than a dozen individuals. He also observed one larger aggregation near YaoundE,
Cameroun.
In Cameroun, B. tuberosus breeds during the dry season and at the beginning
of the rainy season fi principally January-March (Amiet, 1976).
Comparisons with other species. The mating call of B. tuberosus is as distinc-
tive as its morphology. Table 10 compares the acoustic structure to those of twenty
African species including all species whose mating calls are known from west and
central Africa and all members of the B. regularis Complex (Tandy & Tandy, 1976).
See also data in Tandy et al., 1976; Largen et al., 1978; and Tandy et al., 1982 and
1985. The call of B. tuberosus appears to differ diagnostically (ranges of variation do
not overlap in data available) in at least 3 characters from all twenty species. This
comparison must be considered tentative because of the small sample size (2)
available for B. tuberosus, but the comparisons are probably valid because of the
much larger data sets available for most of the other species.
Only B. gracilipes has been observed calling at the same time and locality as
B. tuberosus, although B. camerunensis has been observed breeding nearby. Other
species of the central African rain forests that might breed in the same areas include
B. latifrons, B. maculatus and B. superciliaris. The latter is thought to lack a mating
call (Amiet, 1976).
The call of B. tuberosus differs in five of six apparently homologous characters
from B. gracilipes, B. villiersi and B. perreti and it also differs qualitatively from
those species. The mating calls of the above three species are first order sequences of
complex pulse trains whereas that of B. tuberosus 1s just one complex pulse train.
The voice of B. camerunensis differs in all six call characters compared. The
PPR is slower in B. camerunensis, the NPP/PT and PT DUR two to three times
greater and PTR is one third to one fourth its rate in B. tuberosus.
The. mating call of B. latifrons is different in all six characters. PT DUR in 2.
tuberosus is about one fifth, PPR is more than three times faster and PTR is two to
four times faster than in B. latifrons.
The voice of B. maculatus differs in four characteristics. PPR and PT DUR are
about twice, NPP/PT four times and PTR up to twice greater than in B. tuberosus.
The call of B. regularis differs in three characters. PPR is less than a third, PT
DUR is about four times greater and PTR less than one half those in B. tuberosus.
These two species are not likely to be sympatric because of different habitat pre-
ferences.
None of the other species in Table 9 is likely to be sympatric with B.
tuberosus. Their calls differ from the latter species in at least three characters.
THE BUFO TUBEROSUS SPECIES GROUP 413
TABLE 9. Apparent differences and similarities between the mating calls of Bufo tuberosus and
19 species of Bufo from west or central Africa and all those known of species of the B.
regularis Complex. * species known from west of the Dahomey Gap. “ species of central
Africa. These comparisons are tentative because of the small number of specimens recorded for
B. tuberosus (n=2). A difference (+) indicates that ranges of variation do not overlap. These
comparisons cannot be tested for statistical significance because of the small sample size for B.
tuberosus and a few of the other species. (1) - species for which number of specimens analyzed
is less than 10. For data on other species, see Table 1 in Tandy et al. (1976), Table 2 in Largen
et al. (1978), Table 1 in Tandy er al. (1982), and Table 14 in Tandy & Feener (1985). (2)
Comparison is between homologous characters rather than those of the same grade of physical
compexity (See Tandy & Tandy, 1976).
Species compared PPR NPP/PT PTDUR PTR LOW EMP DOM Number of characters
Different Similar
+ (2) + (2)
+ (2) + (2)
B. gracilipes* +
villiersi “(1) +
. regularis** +
garmani =
latifrons (1) +
brauni -
gutturalis +
kisoloensis (1) -
poweri +
rangeri +
camerunensis*" +?
+
+
+
+
+
+
+
+
lea
"9
+
So
+++ + +
+++++ +++ ++.
kerinyagae (1)
xeros*
asmarae
maculatus*"
langanoensis (1)
turkanae (1)
perreti
steindachneri”\(1)
B. pentoni** - Senegal (1)
+
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+
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++++++ +.
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oe ++:
+
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GEOGRAPHIC DISTRIBUTION AND ECOLOGY
Bufo tuberosus is known from fiftyone equatorial lowland rainforest localities
in Cameroun, Guinea Ecuatorial (Bioko and Rio Muni), Gabon, République du
Congo, République Centrafricaine and République Démocratique du Congo between
latitudes 0°5’S and 5°14’N and longitudes 8°33’ and 29°15’E (Fig. 5). These are in an
altitudinal range of 10 to 1266m. See Table 4 for climatic data.
Ecological data in Tables 4 and 10 indicate that B. tuberosus localities differ
diagnostically from those of fifteen of the 25 species compared. B. tuberosus is not
likely to be sympatric with any of those.
Among the remaining ten species, seven occur in central Africa, B. gracilipes,
B. superciliaris, B. latifrons, B. camerunensis, B. maculatus and B. steindachneri. As
noted above, B. tuberosus has been found syntopic only with B. gracilipes. But it is
likely to occur within close proximity of all of these except B. steindachneri. The
latter occurs primarily in marshy sites surrounded by arid savanna.
B. tuberosus is known only from forested localities, so it is not so likely to
occur with B. regularis or B. maculatus which often prefer more open habitats.
414 MILLS TANDY & JEAN-LUC PERRET
The other high forest species, B. gracilipes, B. superciliaris, B. latifrons and B.
camerunensis are most likely to occur with B. tuberosus.
As noted previously, the ecology of B. tuberosus sites is very much like that of
the allopatric B. amieti.
TABLE 10. Apparent differences and similarities between the ecological characteristics of Bufo
tuberosus and species of Bufo from west or central Africa and those of all species of the B.
regularis Complex. * species known from west of the Dahomey Gap. “ species known from
central Africa. Some of these comparisons are tentative because of the small number of known
localities for some species. A difference (+) indicates that ranges of variation do not overlap.
These comparisons have not been tested for statistical significance. (1) - species for which
number of localities is less than 10. For data on other species, see Tables 3 in Tandy et al.
(1976) and Tandy er al. (1982), Table 8 in Tandy er al. (1985), and Table 13 in Tandy &
Teener (1985).
Species compared Elevation Mean Annual Mean Annual Different Similar
Temperature Rainfall
B- Amieti (1) = È = 0 3
B. gracilipes * - 5 è 0 3
B. villiersi * (1) + 3 I 2
B. regularis *’ = 0 3
B. superciliaris ** - = = 0 3
B. garmani - È fu I 2
B. latifrons * (1) = Z Le 0 3
B. togoensis (1) = 2 TE I 2
B. brauni (1) = N È I 2
B. gutturalis = = Fe 1 2
B. kisoloensis * = N x 2 1
B. poweri = È + I 2
B. rangeri = + = | 2
B. camerunensis ** - = 0 3
B. kerinyagae + + - 2 1
B. xeros ** = 2 + I 2
B. asmarae = = + 1 2
B. maculatus ** = 2 È 0 3
B. danielae * (1) - - = 0 3
B. blanfordi = = st | 2
B. langanoensis (1) = a a I 2
B. turkanae (1) = ave SÈ 2 1
B. perreti (1) = is r 0 3
B. steindachneri” = x = 0 3
= 2
pentoni **- Senegal (1) - =
ACKNOWLEDGEMENTS
M. Tandy expresses his appreciation to the late W. F. Blair for providing funds
via National Science Foundation Grant 5406X which supported his fieldwork. He also
thanks his wife Judy for her continued encouragement and support. He thanks his
former wife, J. Hooper for her assistance with field study.
We thank J.-L. Amiet (Université Fédérale du Cameroun) for making available
his recordings of Bufo gracilipes, B. latifrons and B. tuberosus.
We are grateful to the following for providing preserved or living material
from collections in their care: J.-L. Amiet; the late C. M. Bogert, R. Zweifel, C. W.
Myers and G. Foley (New York); J. E. Böhlke, L. Keane and E. V. Malnate
THE BUFO TUBEROSUS SPECIES GROUP 415
(Philadelphia); W. Böhme (Bonn); A. G. C. Grandison and B. Clarke (London); J.
Guibé, E. R. Brygoo, A. Dubois and J. J. Morere (Paris); M. S. Hoogmoed (Leiden);
R. F. Inger and H. Marx (Chicago); M. Lamotte (Paris); A. E. Leviton, R. C. Drewes
and T. Papenfuss (San Francisco); the late R. F. Martin, A. Anderson, D. Canatella
(Austin); C. J. McCoy (Pittsburgh); The late R. Mertens and K. Klemmer (Frankfurt
a. M.); J.-L. Perret and V. Mahnert (Genève); the late G. Peters, P. Beurton and
R. Guenther (Berlin); the late J. A. Peters, R. Crombie, R. Heyer, F. McCullough and
R. Zug (Washington); U. Rahm and C. Unternahrer (Bale); G. Sacarrao and M.
Noqueira (Lisbon); E. Balletto, M. A. Cherchi, the late G. Scortecci and P. Versace
(Genoa); the late R. C. Stebbins and D. B. Wake (Berkeley, California); the late
D. W. Tinkle, R. A. Nussbaum and M. H. Eckhardt (Ann Arbor, Michigan); E. E.
Williams, J. Rosado and P. Kerfoot (Cambridge, Massachusetts) and J. W. Wright
(Los Angeles).
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REVUE SUISSE DE ZOOLOGIE
Tome 107 — Fascicule 2
LANG, Claude. Etat trophique du lac de Morat indiqué par le zoobenthos:
tendance 1980-199 8 vee Ro a RR ANA nn UE
AZPELICUETA, Maria de la Mercedes & José O. GARCIA. A new species of
Astyanax (Characiformes, Characidae) from Uruguay river basin in
Argentina, with remarks on hook presence in Characidae...........
BALLERIO, Alberto. Revision of the genus Ebbrittoniella Martinez (Cole-
optera+Scarabaeoıdea: Ceratocanthidae) Zr anna Need.
LIENHARD, Charles. A new desert psocid from Namibia (Insecta: Psocoptera:
Hroslidaeye ne en Te ee ER
LEISTIKOW, Andreas. Terrestrial Isopoda from Guatemala and Mexico
(Crustacea Oniscidea: Crinocheéta) >
BURCKHARDT, Daniel & Mariängela GUAJARA. Euphalerus clitoriae sp. n., a
new psyllid species from Clitoria fairchildiana (Fabaceae, Papiliono-
ideae), and notes on other Euphalerus spp. (Hemiptera, Psylloidea). . .
KLoss, Kerstin & Wolfgang PFEIFFER. Zur Biologie des “Eingeweide-
fisches” Carapus acus (Brünnich, 1768) (Carapidae, Teleostei), mit
Hinweisen auf eine nicht-parasitische Ernährung. .................
BEDOS, Anne & Louis DEHARVENG. Un nouveau Collembole Neanurinae du
Sud du Vietnam, Blasconura batai sp. n., avec une clé des especes du
genre(CollembolasNeanuridac) è Corr
BAUR, Hannes & Felix AMIET. Die Leucospidae (Hymenoptera: Chalcido-
idea) der Schweiz, mit einem Bestimmungsschliissel und Daten zu den
CUFOpdischenvAnteMm ar o i Eee
TANDY, Mills & Jean-Luc PERRET. The Bufo tuberosus species group with
the description of a new species from the rainforest of Cöte-d’Ivoire. . .
Pages
233-243
245-257
259-275
277-281
283-323
325-334
335-349
351-357
359-388
389-418
REVUE SUISSE DE ZOOLOGIE
Volume 107 — Number 2
LANG, Claude. Trophic state of Lake Morat indicated by the zoobenthos: the
NOS ORI OO So ae 120: ren te I EE
AZPELICUETA, Maria de la Mercedes & José O. GARCIA. A new species of
Astyanax (Characiformes, Characidae) from Uruguay river basin in
Argentina, with remarks on hook presence in Characidae...........
BALLERIO, Alberto. Revision of the genus Ebbrittoniella Martinez (Cole-
optera: Scarabaeoidea:-Ceratocanthidae): EE a O
LIENHARD, Charles. A new desert psocid from Namibia (Insecta: Psocoptera:
Miro Silica) iver rs cc ein ord er ER ETS
LEISTIKOW, Andreas. Terrestrial Isopoda from Guatemala and Mexico
(Crustacea Oniscidea: Crinocheta)........ rn Sea eee
BURCKHARDT, Daniel & Mariängela GUAJARA. Euphalerus clitoriae sp. n., a
new psyllid species from Clitoria fairchildiana (Fabaceae, Papiliono-
ideae), and notes on other Euphalerus spp. (Hemiptera, Psylloidea). . .
Koss, Kerstin & Wolfgang PFEIFFER. On the biology of the pearlfish
Carapus acus (Brünnich, 1768) (Carapidae, Teleostei), with indica-
tions of amon-parasitic nutrition» +2 <= = eres ee RES
BEDOS, Anne & Louis DEHARVENG. A new neanurine springtail from south-
ern Vietnam, Blasconura batai sp. n., with a key to the species of the
genus: (GollembolasNeanuridae)2.. 2.2. RARA de E
BAUR, Hannes & Felix AMIET. The Leucospidae (Hymenoptera: Chalcido-
idea) of Switzerland, with a key and data on the European species. . . .
TANDY, Mills & Jean-Luc PERRET. The Bufo tuberosus species group with
the description of a new species from the rainforest of Côte-d'Ivoire...
Indexed in CURRENT CONTENTS, SCIENCE CITATION INDEX
Pages
233-243
245-257
259-275
277-281
283-323
325-334
335-349
351-357
359-388
389-418
PUBLICATIONS DU MUSEUM D'HISTOIRE NATURELLE DE GENEVE
CATALOGUE DES INVERTEBRES DE LA SUISSE, N° 1-17 (1908-1926) ..... serie Fr. 285.—
(prix des fascicules sur demande)
REVUE DE PALÉOBIOEOGIE mL Nenn. Echange ou par fascicule Fr.
LE RHINOLOPHE (Bulletin du centre d'étude des chauves-souris)
COLLEMBOLENFAUNA EUROPAS
von H. Gisin, 312 Seiten, 554 Abbildungen, 1960 (Nachdruck, 1984) ............. Fr.
THE EUROPEAN PROTURA THEIR TAXONOMY, ECOLOGY AND
DISTRIBUTION WITH KEYS FOR DETERMINATION
are par fascicule Fr.
by J. NOSEK, 346 pages, 111 figures in text, 1973.............................. Fr.
CLASSIFICATION OF THE DIPLOPODA
PARRIChAr dE HOREMAN; 237) pases) 1979 N oe ee Fr.
LES OISEAUX NICHEURS DU CANTON DE GENEVE
par P. GEROUDET, C. GUEX et M. MAIRE
351 pages, nombreuses cartes et figures, 1983 ................................. Fr.
CATALOGUE COMMENTÉ DES TYPES D'ECHINODERMES ACTUELS
CONSERVES DANS LES COLLECTIONS NATIONALES SUISSES,
SUIVI D'UNE NOTICE SUR LA CONTRIBUTION DE LOUIS AGASSIZ
A LA CONNAISSANCE DES ECHINODERMES ACTUELS
par Michel JANGOUX, 67 pages, 11 planches, 1985 ............................. Fr.
RADULAS DE GASTÉROPODES LITTORAUX DE LA MANCHE
(COTENTIN-BAIE DE SEINE, FRANCE)
par Y. FINET, J. WUEST et K. MAREDA, 62 pages, 1991 .......................... Fr.
GASTROPODS OF THE CHANNEL AND ATLANTIC OCEAN;
SHELLS AND RADULAS
by NOEINET TAWUESR and 'KMAREDA 1992 MER PEN RE ee: Fr.
O. SCHMIDT SPONGE CATALOGUE
par R. DESQUEYROUX-FAUNDEZ & S.M. STONE, 190 pages, 49 plates, 1992 .......... Fr.
ATLAS DE RÉPARTITION DES AMPHIBIENS
ET REPTILES DU CANTON DE GENEVE
par A. KELLER, V. AELLEN et V. MAHNERT, 48 pages, 1993 ...................... Fr.
THE MARINE MOLLUSKS OF THE GALAPAGOS ISLANDS:
A DOCUMENTED FAUNAL LIST
PARMVES EINER LS 0ipases 100 RECON I EE Fr.
NOTICE SUR LES COLLECTIONS MALACOLOGIQUES
DU MUSEUM D'HISTOIRE NATURELLE DE GENEVE
par Jean-Claude CAILLIEZ, 49 pages, 1995 .................................... Fr.
PROCEEDINGS OF THE XIIIth INTERNATIONAL CONGRESS
OF ARACHNOLOGY, Geneva 1995 (ed. Volker MAHNERT), 720 pages, 1996 ....... Fr
CATALOGUE OF THE SCAPHIDIINAE (COLEOPTERA: STAPHYLINIDAE)
(Instrumenta Biodiversitatis I) par Ivan LOBL, 192 pages, 1997.................... Fr.
CATALOGUE SYNONYMIQUE ET GEOGRAPHIQUE DES SYRPHIDAE (DIPTERA)
DE LA REGION AFROTROPICALE
(Instrumenta Biodiversitatis II) par Henri G. DIRICKX, 188 pages, 1998
A REVISION OF THE CORYLOPHIDAE (COLEOPTERA) OF THE
WEST PALAEARCTIC REGION
(Instrumenta Biodiversitatis III) par Stanley BOWESTEAD, 204 pages, 1999 ............ Fr.
THE HERPETOFAUNA OF SOUTHERN YEMEN AND THE
SOKOTRA ARCHIPELAGO
(Instrumenta Biodiversitatis IV) par Beat SCHATTI & Alain DESVOIGNES,
WF Br pases 1999 Ar En A RE LEER Fr
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Volume 107 - Number 2 - 2000
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